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Research ArticleSugarcane Landraces of Ethiopia Germplasm Collection andAnalysis of Regional Diversity and Distribution
Esayas Tena Gashaw 1 FirewMekbib2 and Amsalu Ayana3
1Ethiopian Sugar Corporation Research and Development Eastern Shewa Zone PO Box 15 Wonji Ethiopia2HaramayaUniversity College of Agriculture and Environmental Sciences School of Plant Sciences PO Box 138 Dire Dawa Ethiopia3Integrated Seed Sector Development Ethiopia Program Addis Ababa Ethiopia
Correspondence should be addressed to Esayas Tena Gashaw esutena11gmailcom
Received 8 May 2018 Revised 6 July 2018 Accepted 19 July 2018 Published 14 August 2018
Academic Editor Clifford Gold
Copyright copy 2018 Esayas Tena Gashaw et al This is an open access article distributed under the Creative Commons AttributionLicense which permits unrestricted use distribution and reproduction in any medium provided the original work is properlycited
Sugarcane has been cultivated by smallholder farmers since 16th century in Ethiopia and preceded the commercial productionHowever as far as this study is concerned no exploration and collection have been conducted to know the landraces and study theregional diversity of the crop Therefore the objectives of this study were to collect native sugarcane landraces in Ethiopia and toassess phenotypic diversity and analyze regional distribution among landraces collected from different geographical regions Morethan 300 sugarcane genotypes were collected The landraces were analyzed for 21 quantitative stalk and juice quality charactersand 16 qualitative characters Phenotypic diversity among landraces was high as expressed by the large range of variation for meanquantitative traits and the high (080) ShannonndashWeaver diversity index Our results provided experimental evidence on occurrenceof geographical variation and significant within-region variation where it was high in the regions of Amhara Benshangul-Gumzand SNNPR Wide variability of agronomically important characters in sugarcane such as millable stalk count at harvest singlecane weight and plant height was observed among regions These characters also demonstrated high correlation with cane andsugar yield and the altitude of the collection sites Therefore breeders can utilize accessions of regions showing variability forthese characters in selection programs and to design breeding strategies to produce varieties with best commercial merits Thepresent study contributes to updating sugarcane descriptors adopted from USDA-ARS as well as Bioversity passport data forthe future collection and evaluation The paper discussed insinuation of the results with regard to plant breeding germplasmcollection and conservation as well as the plausible sources for the wide range of variation observed This is the first study toreport landrace sugarcane genetic resources in Ethiopia and information on geographical pattern of variation in Ethiopian localsugarcane germplasm
1 Introduction
Sugarcane plays a significant role in the Ethiopian socioecon-omy Sugar and its byproduct are used for local consumptionand export The industry created job opportunity for a largenumber of people Today in the country sugar consumptionoutstrips its production The per capita sugar consumptionin Ethiopia is very low (5-6 kg) which is even below theAfrican standard (15 kg) while the world average per capitaconsumption is 21 kg in 2016 The commercial sugarcanesector in Ethiopia commenced since 1951 Sugar Corporationof Ethiopia currently administers six sugar factories namely
Wonji-Shoa Metahara Finchaa Tendaho Arjo-dedessa andKessem and nine sugar development projects at Kuraz TanaBeles and Welkayit Sugarcane plantations are expandingwith current area coverage of 98986 hectares and productionof 400000 tons of sugar and 25388m3 of ethanol perannum The new sugar factories planned to have ethanoland cogeneration facilities thereby increasing the productionof sugar and coproducts Accordingly when all projects arecompleted the annual sugar production will be boosted to39-417 million tons ethanol production will be 181 millionlitres and the factories contribute 709 Mega Watt electricpower to the national grid This is 118 of sugar production
HindawiAdvances in AgricultureVolume 2018 Article ID 7920724 18 pageshttpsdoiorg10115520187920724
2 Advances in Agriculture
by the leading sugar producer Brazil with a total amount of353 million tons produced in 201617 [1] Similarly Brazilianethanol production reached 3023 billion litres in 201516
Though commercial sugarcane production has a historyof six decades sugarcane had been cultivated in Ethiopiasince 16th century According to the report by central statisticsagency (CSA) currently sugarcane is produced in about3123681ha with 156506000 holdings in different parts ofthe country [2] But the production is not usually usedfor industrial purposes It is noticeably used for makingconfectioneries household consumption (chewing) sellingfor immediate cash and feeding livestock In some areassugarcane is used to prepare local beverage called ldquoKaribordquomainly preferred byMuslim communities while in others theleaves are used for thatching and as firewood [3] Howeverthe potential of this sector is not well explored and has notbeen given due consideration Furthermore no explorationand germplasm collection have been done to represent andpreserve local landraces
The sugar industry of Ethiopia is so far dependent onintroduction of exotic varietieswhich are not suitably adaptedto various agroecologies and local growing conditions Inlight of the rapidly increasing commercial sugarcane planta-tion areas in the country the demand for improved varietiesthat suit various agroecologies is increasing Under suchsituations there will be a continuous demand for broadgenetic base sugarcane varieties that are high yielding andstable under abiotic and biotic stressesTherefore the industryis currently launching breeding program which is long over-due to produce its own improved varieties The developmentof high yielding and stable varieties requires a continuoussupply of new germplasm as a source of desirable genesandor gene complexes The primary sources of such genesare landraces introductions weedy and wild relatives of cropplants [4 5] The availability of such germplasm requiresthe identification of areas of diversity of various charactersof agronomic importance especially in the local landracesgrowing within the variable agroecologies of Ethiopia There-fore germplasm collection and conservation and the studyof genetic diversity of Ethiopian sugarcane landraces areworthwhile since this can broaden the genetic base andprovide locally adapted genes for improvement of the crop
In spite of the great importance of Ethiopian sugarcanelandraces for the germplasm genetic base improvement andutilization in the breeding program no effort has beenmade so far to collect and preserve this genetic wealthFurthermore study of the variation and assessment of extentand geographical pattern of distribution of this landracesis lacking For effective utilization of germplasm in plantbreeding programmes the information on the extent andpatterns of distribution of genetic variation of a crop species isvery essential [6ndash8]This also canhelp in devising appropriatesampling procedures for germplasm collection and conser-vation and obtaining core collection for efficient germplasmmanagement [9 10]
The objectives of this study were to collect sugarcanelandraces in Ethiopia and to assess phenotypic diversity andanalyze regional distribution among landraces collected fromdifferent geographical areas
2 Materials and Methods
21 Germplasm Collection
211 Sampling Technique Sugarcane germplasm were col-lected during 201011 all across Ethiopia in the regionalstates of Amhara (07102010-09052011) Oromia (26082010-08072011) Southern Nations Nationalities andPeoples Region (SNNPR) (10082010-09122010) Tigray(21042011-09052011) Benshangul-Gumz (20122010-15022011) Gambella (04112010-28112010) Somali (29062011-26072011) and Harari (02062011-23062011)(Figure 1 Supplemental Table 1) Collection was made fromhomesteads farmersrsquo fields and local markets Germplasmwas collected using stratified random sampling techniquesampling areas are shown in Figure 1 In each region all zoneswere sampled Two to four districts (locally referred to asldquoWeredasrdquo) were selected from each zone From each district(depending on size a district contains several localities orsubdistricts) 2-5 subdistricts (locally referred as ldquoKebelesrdquo)or peasant associations (PAs) were selected The districts andsubdistricts were selected based on long agricultural historyand relatively wide areas allocated to sugarcane productionMoreover purposive sampling was also employed based oninformation supplied by key informants on the unique andquality sugarcane types grown in these areas In the selectedsubdistricts sugarcane clones were collected following themethods proposed in [11] Each distinct morphotype in avillage was randomly sampled Information on the sampledsugarcane germplasm was recorded and passport data wascollected following the method of Bioversity International[12] Moreover juice sample was taken from the bottommiddle and top part of the stalk of each clone and meanpercent brix reading was recorded using hand refrac-tometer
212Determination of the Physical andChemical Properties ofSoils in SugarcaneGermplasmCollectionAreas Todeterminethe predominant physical and chemical properties and thefertility status of soils under sugarcane production sampleswere collected and analyzed across germplasm collectionareas Georeferencing (latitude and longitude) of the studysites were made with a Garmin GPS In every germplasmsampling area a composite soil sample was taken between0ndash30 and 30ndash60 cm depths Soil samples were analyzed fororganic carbon total nitrogen (N) soil pH soil electricalconductivity (EC) available phosphorus (P) and availablepotassium (K) contents using standard procedures [13] SoilpH was measured potentiometrically using a digital pHmeter (Jenway Model-3320 GransmoreGeeen) Soil EC wasmeasured using digital conductivity meter (Jenway Model-4310 GransmoreGeeen) Organic carbon was determinedfollowing the wet digestion method described in [14] Total Nwas determined using the Kjeldahal procedure [13] Availablephosphorus was determined using the Olsen method [15]and Bray II method [16] for acidic soils Available K wasmeasured by flame photometry using the sodium acetateextractant method at pH 48 [17] Soil texture was determinedby hydrometer method [17]
Advances in Agriculture 3
1700000
1500000
1300000
1100000
900000
700000
500000
300000
1700000
1500000
1300000
1100000
900000
700000
500000
300000
minus300000 100000 500000 900000 1300000 1700000
minus300000 100000 500000 900000 1300000 1700000
Sampling Site Location Map
Figure 1 A map of Ethiopia showing sugarcane germplasm collection sites (black circles)
Table 1 Location information of the test sites
Location Latitude Longitude Altitude (masl)119886 Rainfall (mm) Temperature (∘C)Minimum Maximum
Wonji 8∘311015840 N 39∘121015840 E 1550 800 153∘C 269∘CMetehara 8∘511015840 N 39∘ 521015840 E 950 554 175∘C 326∘C119886masl = meters above sea levelSource meteorological Stations of respective locations
22 Diversity Study
221 Plant Materials A total of 211 sugarcane (Saccharumspp) accessions consisting of 196 landraces (SupplementalTable 3) and 15 introduced commercial varieties including2 standard varieties (Supplemental Table 4) were used forthis study The landraces represent collections from allgeographical regions across Ethiopia maintained at fieldconservation garden at Wonji and Metehara Sugar EstatesSugar Corporation of Ethiopia Sampling of the landraces wasmade on representation basis stratified systematic samplingmethod to a given range of geographic area altitudinalranges and a range of morphological traits The introducedmaterials were commercial varieties under production indifferent estates where the two were standard varieties
The 196 landraces were collections from the followingregions of Ethiopia Amhara (47) Benshangul-Gumz (10)Gambella (3) Harari (2) Oromiya (65) SNNPR (59) Somali(3) and Tigray (7) (Figure 1) The altitude of the collectionsites for the landraces used in this study ranged from 454to 2687 meters above sea level representing the distributionof the crop in Ethiopia The introduced materials were fromBarbados (6) Cuba (1) India (5) Mexico (1) and SouthAfrica (2)
222Methods The plant materials were grown at Wonji andMetehara Sugar Estates of Sugar Corporation of Ethiopiaduring the 201213 growing season Details of the plantinglocations are shown in Table 1 Each accession was grown ina single row plot of 5m long and 145m between rows and20 cm between plants within a row with two replications inrandomized complete block design
Uniform crop management practices were applied asrecommended for the areas Urea was applied 25 monthsafter planting at a rate of 200 kgsdothaminus1 atWonji and 400 kgsdothaminus1at Metehara
Accessions from regions with sample size less than 12were included in adjacent regions to reduce experimentalerror due to small sample size Hence the two Harari andthree Somali accessions were included in Tigray and thethree Gambella accessions were included in Benshangul-GumzThis reduced the 8 regions of Ethiopia fromwhich thelandraces were originally drawn to five With the introducedmaterials included 6 regions of origin were used in thestatistical analyses
Data on 17 quantitative stalk characters was recordednamely sprout count 1 and 2 months after planting(SPC1MAP and SPC2MAP) tiller counts 4 and 5month afterplanting (TC4MAP and TC5MAP) stalk count 10 months
4 Advances in Agriculture
after planting (STC10MAP) hand refractometer brix reading10 months after planting (HRBrix10MAP) millable stalkcount per hectare (MSCHA) single cane weight (SCW)number of internode (NOI) internode length (IL) stalkheight (SH) stalk diameter (SD) leaf length (LL) leaf width(LW) leaf area (LA) cane yield quintal per hectare (CYHA)and sugar yield quintal per hectare (SY) Data on 4 juicequality parameters ie brix percent (Brix) pol percent(pol ) purity percent (purity ) and sugar percent (SR) was also recorded For every accession ten plants wereused for recording data for quantitative characters whichwere recorded on plot basis Count data and cane yield wererecorded considering all cane stalks from the whole plot Forquantitative leaf characteristics measurement a proceduredeveloped in [18] was used
To categorize each accession morphologically sugarcanedescriptors adopted fromUSDA-ARSwere employed (GRIN2004) Data on 16 qualitative traits was recorded namelypresence or absence of bud cushion (BUDCUSHION) rel-ative degree of bud extension (BUDEXTEND) relative budshape (BUDSHAPE) relative shape of dewlap (DEWLAP-SHAP) type of outer auricle (AURICLEOUT) presence orabsence of stalk corky cracks (STALKCORKC) presenceor absence of stalk corky patches (STALKCORKP) relativeshape of ligule (LIGSHAPE) presence or absence of stalkgrowth cracks (STALKCRACK) presence or absence of budgroove (BUDGROOVE) relative plant erectness (ERECT)relative degree of internode alignment (INALIGN) relativeinternode shape (INSHAPE) colour of the leaves (LEAF-COLOR) colour of the exposed rind (RINDCOLE) andcanopy structure (CANOPY) Each accession was scored forthe most frequent character-state Leaf colour and colourof the exposed rind were examined and scored using theMunsell colour chart [19]
223 Statistical Analyses First analysis of variance wasmadefor the 21 characters for each location Homogeneity ofthe error variances among the locations was assessed in[20] F-max method for each of the 21 characters The testestablished the homogeneity of the error variances for allcharacters except TC4MAP For TC4MAP logarithmic datatransformation which is recommendable for continuousdata was used to homogenize the error variance Thendata of all the characters were subjected to test location-accession-related analysis of variance to determine effects oftest sites and genotype X environment (G X E) interactionThe results showed the nonsignificance of effects of testsites for SPC1MAP IL LW Pol sugar percent (SR) andthe nonsignificance of G X E interactions for SPC2MAPSCW Brix Pol and Brix10MAP For the other charactersboth test site effects and G X E interactions were significant(Plt005)
Analysis of variance was made for 21 quantitative char-acters following the procedure used in [21 22] The meansquares of the regions were tested against pooled meansquares of accessions within regions The pooled meanssquares for accessions within regions of origin and the meansquares of accessions within each region were tested against
the pooled within-region error mean squares Means rangesfor means and percent coefficients of variation for all thecharacters were computed for each region of origin and forthe entire data The regional means were compared usingDuncanrsquos multiple range testing Correlations between thecharacters were computed at three levels First correlationsof the characters were assessed based on the 211 accessionmeans Then interregion correlation was computed usingthe means of characters for each region Finally a series ofintraregion correlation coefficient matrices were obtained foreach region using the accession means from that region forthe characters
For qualitative characters phenotypic frequency distri-butions were worked out for all the sample germplasm andlocations The ShannonndashWeaver diversity index (H1015840) wascomputed using the phenotypic frequencies to assess thephenotypic diversity for each character for all accessions TheShannonndashWeaver diversity index as described in [23] is givenas
1198671015840 = minus119899
sum119894=1
119901119894log119890(119901119894) (1)
where pi is the proportion of accessions in the 119894th class of ann-class character and n is the number of phenotypic classesof traits Each 1198671015840 value was divided by its maximum value(logen) and normalized in order to keep the values between0 and 1 By pooling various characters across the regions theadditive properties of 1198671015840 were used to evaluate diversity ofregions and characters within the population The averagediversity index (H1015840) over n traits was estimated as H1015840 =sumH1015840n
3 Results and Discussion
31 Collection of Sugarcane Germplasm In Ethiopia thehistory of sugarcane cultivation by smallholder farmers pre-ceded that of commercial cultivation As documented in thehistory of a monastery in Northern Ethiopia sugarcane hasbeen grown in the country since the 16th century [2] In thisstudy local sugarcane germplasm exploration and collectionwere conducted all across Ethiopia in the regional statesMore than 300 local sugarcane genotypes were collectedduring 20102011 and passport data of the genotypes ispresented in Supplemental Table 1 The collected germplasmwere planted at five locations across the country with respectto their collection area These are Wondogenet AgriculturalResearchCenter JimmaAgricultural ResearchCenterMechaWereda Agricultural Bureau nursery field at Picolo AbaySirinka Agricultural Research Center (at Kobo Subcenter)and research field of Haramaya University at Diredawa(Tonny Farm) The clones have been monitored during the201112 season for symptoms of major diseases and insectsThese clones were transferred to commercial sugarcane plan-tation estates at Wonji and Metehara for further selectionand maintenance No major diseases and insect pests weredetected in all the germplasm collected except sometimestermites and borer were observed (Supplemental Table 2)
Advances in Agriculture 5
Table 2 Mean soil physical and chemical properties of sugarcane fields in the surveyed regions
Regions Soil chemical properties Texture ()PH EC(dsm) OC N P (ppm) K(ppm) Sand Silt Clay
Most of the sampled germplasm had acceptable levels of juicerefractometer readings (expressed in degree brix) relative tothe history of the ages of the samples (Supplemental Table 1)The number of harvestablesaleable cane stalks obtained inplant cane and ratoon crops and the number of harvestableratoon crops showed variation region to region and zone tozone and sometimes district (ldquoweredardquo)-wise There is alsovariation in maturity time of the clones both in plant caneand in ratoon crops (Supplemental Table 2)These are mainlydepending on the variety soil condition cultural practicesand climatic conditions
The genotypes collected from the respective zones inthe study regions were recorded by their local namesSometimes similar varieties bear different names at differentlocalities and vice versa Farmers mentioned a broad rangeof sugarcane landraces that had been grown in the areasand maintained for generations Some of the landraces werecommonly recognized by most farmers within and acrosszones of the respective regions whereas some were rarevarieties known only by few farmers The large number oflandraces observed during the current study demonstratesthe existence of diverse genetic resources of sugarcane inEthiopia The diversity could have evolved presumably due todiverse climatic conditions low input management systemshigh pests and disease pressure and continuous selectionby farmers Smallholder farmers face diverse environmentstresses and have multiple production objectives that affectselection of genotypes [24] There were no formally releasedimproved sugarcane varieties grown by farmers in the studyregions except in few places This was mainly observedin SNNPR at Wolayta and Kembata-Tembaro zones Thesevarieties have been informally introduced to these areas byseasonal workers employed at the sugar estates especially atWonji and Metehara
The collected landraces would serve to broaden thegenetic base of the available sugarcane germplasm Thesugar industry in Ethiopia is currently establishing sugarcanebreeding programTherefore collection and efficient charac-terization of germplasm are vital to strengthen the breedingprogram
311 Physical and Chemical Properties of Soils of Sugar-cane Germplasm Collection Areas Table 2 summarizes thephysical and chemical properties of soils in the sugarcane
production fields of surveyed areas Sugarcane productionby smallholder farmers in the surveyed areas is conductedacross the seasons where harvesting and planting are done aslong as there is available moisture Therefore the soil samplesrepresent the sugarcane production season and the altitudinalranges of the respective areas
Generally the organic carbon content of soils in sugarcaneproducing areas across the country is low to very low (2-4lt2) Amhara Gambella SNNP Somali and Tigray Regionsare in a very low category (lt2) Similarly across regions thetotal N content is low except Harari and Oromia regionswhere the average total N is medium This problem shouldbe addressed either with augmenting the soil with inorganicfertilizers or through appropriate management practices likecrop residue management Available phosphorus is high inAmhara Gambella and SNNP Regions whereas moderatevalues were observed in Harari Oromia and Tigray RegionsGenerally the level of phosphorus in soils of sugarcaneproducing areas in these regions is adequate In Benshangul-Gumz and Somali Regions low level of phosphorus wasrecorded which should be addressed with due attentionAvailable K is sufficient for normal sugarcane growth in soilsof all regions Relatively lower values of K were recorded insoils of Harari and Tigray Regions It should be noted thatorganic carbon EC PH and variability of P andK are affectedby rainfall and temperature that are seasonal and the valuesrecorded refer to the seasonwhen the soil samples were takenGenerally Ethiopian soils are deficient in various essentialnutrients like boron nitrogen phosphorus potassium sulfurzinc and copper although severity differs from region toregion [25] Particularly the loss of P and N resulting fromthe use of dung and crop residues for fuel is demonstrated tobe equivalent to the total amount of commercial fertilizer use
Across regions in Ethiopia generally clay type of soildominates followed by silt except in Gambella Region wherethe dominant soil texture was sand Soil management optionsrelated to clay soil may address soil related problems insmallholder sugarcane growing areas across Ethiopia
32 Diversity of Sugarcane Landraces
321 Quantitative Characters
Univariate Statistics Analysis of variance depicted highlysignificant differences (p lt 001) between accessions pooled
6 Advances in Agriculture
over the regions for the 21 characters of the 211 sugarcaneaccessions studied and significant difference for the 6 regionsof origin for 6 characters (Table 3) The results suggestedthe occurrence of significant phenotypic variation betweenthe accessions as a whole Significant variations of differentsugarcane stalk and juice quality characters was also reportedin similar studies conducted elsewhere [26 27] Region-wise partitioning of the variance indicated significant within-region differences (p lt 005) among the populations withinalmost all regions for the characters NOI IL SH SD LLLW LA Brix Pol Purity SR MSCHA SCW CYHAand SY for 8 characters within Tigray for TC4MAP andTC5MAP within Oromiya and SNNPR Brix10MAP showedno significant difference within all regions
In general within-region variation was greater for stalkdiameter single cane weight millable stalk count and caneand sugar yield than for other characters for all the regionsAssuming that a significant portion of the phenotypic vari-ation is genetic it would be possible to make selection forany of this group of characters within a particular regionIt was apparent that the variance between accessions pooledover regions was greater than between regions Therefore inorder to sample the variation effectively it would be necessaryto sample the variable populations from different localitiesin a region The study in [28] on genetic diversity amongmain land USA sugarcane cultivars showed high geneticdiversitywithin populations of different regions than betweenpopulations
Duncanrsquos multiple range testing for regional means for allthe characters is shown in Table 4 Relatively much differenti-ationwas observed for stemdiameter and juice purity percentcompared to other parameters Important yield componentslike MSCHA SCW SH and SD have shown higher valuesin Tigray SNNPR and Oromiya regions High cane andsugar yield were recorded for accessions in Tigray regionfollowed by SNNPR Oromiya and introduction which werestatistically at par Statistically similar lower values of caneand sugar yield were recorded in Amhara and Benshangulcollections
Higher value of sucrose percent in cane was also observedfor accessions from Tigray and the lowest value was inAmhara accessions while other regions were statisticallysimilar Generally for most of the characters high valuewas obtained from accessions in Tigray region followed bySNNPROromiya and introduction Therefore the materialsfrom these regions can be used for selection of cane and sugaryield per se and in breeding program The local accessions inmany of the regions showed superiority over the introducedcommercial varieties currently in production for most of thecharacters including cane and sugar yield (Table 4)This maybe due to the adaptation and coevolution of the materialswith different stresses for long period of time in the countryrsquosagroecology These locally adapted genes conferring yieldadvantage could be harnessed through strategic crossing andselection for improvement of the crop Superiority of locallandraces over introduced varieties was reported [29 30]
The range of variation of the accession means establishedwide variation between the regions and the accessions withinthe regions for the characters studied (Table 5) Accordingly
the maximum score was 41 times the minimum for sproutcount one month after planting 32 for sprout count twomonths after planting 25 for tiller count four months afterplanting 22 for sugar yield 20 for cane yield 12 for millablestalk count 8 for stalk count 10 months after planting 7for single cane weight 6 for tiller count five months afterplanting and 2 for other yield components and sugar qual-ity parameters except hand refractometer brix reading tenmonths after planting and juice purity percent which scored1 The same trend was observed between accessions withina particular region However accessions within AmharaBenshangul-Gumz and introduction showed wider ranges ofvariation than those accessions within other regions for themajority of the characters The wide range of geographic andclimatic features of the regions in Ethiopia and the originaldifferentiation and further farmers selection for productionniches and uses for hundreds of years may have resultedin the accessions to possess a tremendously high degree ofmorphological variation The reason for introduced varietiesshowing high variation also could be due to broader geo-graphic spectrum fromwhere theywere initially acquired anddifference in their parental source and genealogical historyIt was demonstrated even with these limited numbers ofsamples (15 genotypes) that introduction of genetic materialshas practically broadened the genetic base of the sugarcanegermplasm for the breeding program
Coefficient of variation measured as the ratio of standarddeviation to the corresponding overall mean and expressedas percentage is useful to compare different charactersmeasured in different units It is also useful to compare samecharacter in different groups of populations with differentsample size mean and variance or different characters indifferent populations In the current study high coefficientsof variation were observed between regions and within eachregion for sprout as well as tiller counts in the monthsthe data was recorded stalk count 10 months after plant-ing and millable stalk count at harvest cane and sugaryield single cane weight and leaf area (Table 6) Thoughaccessions from a particular region were more variable fora specific character compared to other regions accessionsfromAmhara Benshangul-Gumz introduction and SNNPRwere more variable than accessions from other regions Thisresult demonstrated the tremendous variability of sugar-cane germplasm from these regions The accessions fromTigray and Oromiya had low coefficients of variation formany characters indicating relatively high within-regionuniformity The high coefficients of variation observed formost of the characters agreed well with those reportedin [31] for indigenous sugarcane of Brazil Similar resultswere also reported in sorghum [32] The different levels ofregional variability of a particular character could be due todifferences in forces of selection andor differences in theintensity of a particular selecting force It could also be dueto random effect and reduced sample parents of parentalmaterials moved differently by various human populationmigration and the interaction of these with Darwinianforces
Results of this study suggested the presence of wide rangeof variations of characters determined by univariate statistics
This is in agreement with previous studies on sugarcanegermplasm [33ndash35]
This is the first study to report information on geo-graphical pattern of variation in Ethiopian local sugarcanegermplasm which has been lacking so far Our results pro-vided experimental evidence on occurrence of geographicalvariation and significant within-region variation where it washigh in the regions such as Amhara Benshangul-Gumz andSNNPR
The overall patterns of similarity or difference betweenregions seemed to depend on environmental factors such asrainfall temperature length of growing season and altitudeWide variability of millable stalk count at harvest singlecane weight and plant height was observed among regionswhere these characters demonstrated high correlation withcane and sugar yield (Table 7) Therefore breeders can utilizeaccessions of regions showing variability for these charactersin selection programs and to design breeding strategies toproduce varieties with best commercial merits
Bivariate Statistics Correlation coefficients worked out onthe 21 quantitative characters are shown for the entire data(Table 7) between regions (Table 8) and within regions(data not shown) Results of the association studies for theentire data showed that cane and sugar yield had highlysignificant positive correlation with most of the quantitativestalk characters and juice quality parameters However theassociation of tiller counts 4 and 5 months after plantingstalk count 10 months after planting millable stalk countat harvest single cane weight and plant height with caneand sugar yield was stronger suggesting these componentcharacters asmain contributing factor to cane and sugar yieldIn fact sugar yield is the product of cane yield and sucrose per-cent cane surely being influenced with components affectingcane yield Similar trend was reported by earlier sugarcaneworkers [36 37] In sugarcane the cane and sugar yields areconsidered to be the complex characters The information onthe phenotypic interrelationship of cane yield and sugar yieldwith their component character per se would be of immensehelp to sugarcane breeders
Stem diameter had significant positive association withboth cane and sugar yield indicating the significance ofthis trait in improving both cane and sugar yields Similarreports have beenmade by earlier sugarcane workers [37 38]Stalk diameter was also positively and significantly correlatedwith plant height It is one of the traits that are related tolodging resistance [39] and the positive associations withheight would help in reducing the chance of lodging as heightincreases Among the physiological attributes included in thestudy leaf area had highly significant positive associationwith cane yield Similar observation was also made in [37]Sucrose percent which is an important juice quality attributehad highly significant positive association with sugar yieldand simultaneously cane yield also had highly significantassociation with sugar yield Based on the magnitude ofcorrelation coefficient values it can be said that cane yield ismuch more important than sucrose percent in determiningthe sugar yield and the same has also been reported by othersugarcane workers [37 40] Similar to observations made in
[37 41] results of the current study showed that all the qualityparameters like sucrose percent pol percent brix percentand purity percent were significantly correlated in positivedirection This indicated any of these juice quality traitscould be considered for selection leading to the simultaneousimprovement in the remaining quality traits
Ten of the 21 characters also showed significant positivecorrelations with altitude of the collection sites (Table 7) Asdescribed in [42] ecological characteristics have influencedthe genotypic constitution of landraces during domesticationand hence a relationship exists between the agroecology inthe collection site and the morphological characteristics ofthe landraces Thus positive correlation between collectionsite variables and plant characteristics would imply that thevariation between accessionsmay be related to agroecologicalvariations among the collection sites [43]
The correlation coefficient between number of internodesand altitude was negative and significant indicating thatother environmental factors (other than altitude) andornonenvironmental factorsmight account for the variation forthis particular character Since temperature decreases withan increase of altitude in Ethiopia [44 45] it is more likelythat temperature has exerted strong selection pressure innumber of internodes in the genotypes Information on therelationships between environmental factors of the collect-ing sites and morphophysiological variation of germplasmcould enhance the understanding of evolutionary adaptivepatterns which could assist germplasm collectors and users[46]
Character associations between region and within regionalso followed similar fashion like that of the entire dataNumber of sprout tiller and stalk counts and sugar qualityparameters showed significant positive correlations with caneand sugar yield
Though not significant and strong the correlation of leaflength and width and leaf area (leaf area = leaf length Xleaf width X 075) with cane and sugar yield was negativein the interregion correlations This kind of relationship wasalso exhibited between these leaf characteristics and yieldcomponents like tiller and stalks counts plant height andsingle cane weight and stalk diameter and sugar qualityparameters This may have been attributed to the variationof temperature with altitude and other environmental factorsacross the regions resulting in such type of relationshipSuch relationship also calls for the need to have enough dataon environmental variables of the collection sites such astemperature and rainfall if germplasm collection is requiredtomeet its objectives effectively and efficiently Unfortunatelydata on such useful environmental variables like temperatureand rainfall are lacking in the passport data of the currentEthiopian sugarcane germplasm collections The significanceof environmental factors of the collection sites as veryimportant determinants in structuring morphophysiolog-ical variations was reported in tetraploid wheat [22 4748]
Association of characters among yield its componentsand other economic traits is important for the interpretationof the patterns of variation and making selection in breed-ing program and combining several desirable attributes It
suggests the advantage of a scheme of selection for morethan one character at a time The correlation between char-acters may arise from linkage or from developmental geneticinteractions with orwithout a purely phenotypic componentIt could arise also due to differential phenotypic plasticityof characters themselves Within the limit of experimentalerror and environmental effects high correlation coefficientsbetween characters may show that the characters sharesome common element of genetic control (ie pleiotropylinkage) between genes or else from independently controlledcharacters responding similarly to geographic variation inselection pressures [49] The interregion correlation coef-ficient between the characters measures the consistency oftheir patterns of regional variation while the intraregioncorrelation coefficient measures the association arising fromgenetic factors but is not affected by regional variation[50] Since this study showed significant positive correlationsintraregionally for some character combinations it wouldseem that common genetic control might be playing a role inbringing about correlations between the various charactersIt appeared that different response to regional variationwas playing a greater role than different genetic controlas shown by the many more significant and moderate tohigh correlation coefficients intraregionally than interregion-ally
322 Qualitative Characters The frequency distribution forthe 16 qualitative characters of germplasm samples by regionsin Ethiopia and of the exotic accessions is shown in Supple-mental Table 5 For the purpose of avoiding redundancy andimproving readability the discrete characters are presented bygrouping related traits together
BudCushion andRelativeDegree of BudExtension Evaluationof the presence and absence of bud cushion revealed thatgreat proportion (70) of sugarcane landraces from Ethiopiado not have bud cushion Majority of the sample germplasmin all regions belong to this phenotypic class Approximatelyone-half (53) of the collection have their bud extendedabove the growth ring followed by those (41) having budtouching the growth ring
Relative Bud Shape Among the phenotypic classes for thischaracter tall deltoid was dominant (31) followed by ovate(17) and narrow ovate (12) While local accessions inmost regions had tall deltoid bud shape 50 of the intro-duced accessions belong to two phenotypic classes roundwith central germpore (30) and 20 having pentagonalbud shape Equal proportions (19) of the samples fromBenshangul-Gumz were Ovate and Ovate with emarginatebasal wing
Canopy Structure Relative Plant Erectness Colour of theLeaves and Colour of the Exposed Rind Out of the eightphenotypic classes observed compact tip droopy was themost frequent (35) Open semidroopy and open droopyclasses had fairly equal distribution With regard to relativeplant erectness seventy-five percent of the germplasm werefound to fall in twophenotypic classes erect (41) and almost
erect (34) for the whole region and country Most of theaccessions (90) showed light green or green leaf colourbut 20 from introduced collections were greenish yellowAmong phenotypic classes for colour of the exposed rindbrownish yellow yellow brown and yellowish green were themost frequent with fairly equal distribution
Relative Shape of Dewlap Double crescent had larger fre-quency (22) while squarish descending and flaring typeof dewlap shape had fairly equal proportions Seventeenpercent of germplasm from Tigray region had squarishdeltoid dewlap shape
Relative Degree of Internode Alignment and Internode ShapeForty-four percent of the sample germplasm were slightlyzigzag while there was almost equal frequency for straightand zigzag internode alignment for the entire region andcountry Straight or nearly straight (slightly zigzag) internodealignment of cane stalks is a very important character formechanized farming and postharvest handling in sugarcane[51] Forty percent of the collections were observed havingconcave-convex type of internode shape while 29 wascylindrical and 14 conoidal Other classes had fairly equalproportion
Type of Auricle and Relative Shape of Ligule An equalproportion of the germplasm totally constituting 84 ofthe collection belong to four phenotypic classes namelythose with no auricle and others with transitional and shortand lanceolate auricles Sixteen percent of the collectionsalso showed equal frequency for the remaining classesBenshangul-Gumz and Tigray regions had each equal 13accessions with falcate type of auricle Sixty-seven percentof the collections belong to three ligule shape classes cres-cent with lozenge (27) broad-crescent (21) and linear-crescent (19) The germplasm had almost equal proportionfor the other phenotypic classes of this character Nineteenpercent of the accessions fromBenshangul-Gumz region haddeltoid shape of ligule
Stalk Corky Cracks Corky Patches Growth Cracks and BudGroove Eighty-nine percent of the germplasm across regionshad no stalk corky cracks with only eleven percent exhibitingthis character All the germplasm in Gambella Harari andSomali had no stalk corky cracks On the other hand eighty-one percent of the germplasm for the whole region showedthe presence of stalk corky patches All the local germplasmin Benshangul-Gumz Gambella and Tigray had stalk corkypatches whereas 75 of the germplasm from Harari Region67 from Somali Region and 24 from Oromia Regionhad no stalk corky patches In the entire region 75 ofthe germplasm had no stalk growth cracks with only 25showing this character Sixty-three percent of the germplasmwere found to have bud groove Howevermost of the samplesin Harari (75) Amhara (63) and Oromia (53) Regionshad no bud groove
The results of this study indicated the wide distributionof phenotypic classes for the characters considered whichindicates the existence of different races and combination
Advances in Agriculture 15
of races in local genotypes of different regions in EthiopiaThis diversity of germplasmbased onphenotypicmarkers canhelp sugarcane breeders in identifying the identity of geno-types as well as maintaining genetic diversity Differencesin morphological characters of different sugarcane varietieshave been reported [52] Piscitelli [53] showed importantexomorphological qualitative characters of sugarcane varietythat are not influenced by environmental factors and thus canbe used as a selection tool in any breeding program
The amount of phenotypic diversity estimates based onthe ShannonndashWeaver diversity index (H1015840) are shown inTable 9 The 16 characters differed in their distribution aswell as the amount of variation Individual traits showed adifferent pattern of variation among accessions Estimatesof H1015840 for individual characters varied from 049 for stalkcorky cracks to 100 for bud groove with an overall meanof 080 Most of the characters were highly polymorphicwhereas relative plant erectness and presence or absenceof stalk corky patches scored moderate values The highdiversity values for the characters showed a wide variabilityamong genotypes According to [54] this index is also usedin genetic resource studies as a convenient measure of bothallelic richness and allelic evenness when using geneticdata
The highest diversity computed was on presence orabsence of bud groove with a diversity index of 100indicating that this character is very variable in the localsugarcane germplasm On the other hand the descriptorcomputed with the lowest diversity index value (049) wasthe presence or absence of stalk corky cracks indicatingthat this character is selected in the development of thegenotypes In agreement with the present study [55] workingon S officinarum accessions from the world collection ofsugarcane germplasm obtained a low index of 025 and 035for presence or absence of stalk corky patches and cracksrespectively
Their report also showed high diversity index values forinternode alignment growth cracks internode shape budcushion bud extension bud groove and ligule shape Sim-ilarly [56] experimenting on field collections of sugarcaneaccessions in Philippine reported high ShannonndashWeaverdiversity for stalk growth cracks bud cushion stalk corkypatch stalk corky cracks bud extension and auricle outershape Unlike the present study they have reported high indexfor stalk corky cracksMedium index values were reported forstalk internode shape and low index values for ligule shapeand bud shape High variability of sugarcane varieties fordewlap shape was also reported in [57]
The H1015840 pooled across characters by region ranged from079 in Oromiya to 085 in Tigray with an overall aver-age of 081 (Table 8) Most of the regions showed highShannonndashWeaver diversity index The populations that hadthe highest H1015840 were that of Tigray and introduction The H1015840averaged over regions for different characters was found torange from 044 for presence or absence of stalk corky cracksto 094 for relative degree of internode alignment and budgroove with an overall average of 081 (Table 9)
Assessment of the ShannonndashWeaver index also showedsensitivity to both the number of descriptor classes and the
distribution within phenotypic classes in each region Amongregions the mean values for the ShannonndashWeaver index arenot significantly different but differences were found whenindividual characters were considered These observationssuggest that some regions have high diversity for particulartraits while in other regions selective pressures might havereduced the variation to certain genotypes
4 Conclusions
The large number of landraces observed during the cur-rent study demonstrates the existence of diverse geneticresources of sugarcane in Ethiopia The landraces wouldserve to broaden the genetic base of the available sugarcanegermplasm Generally distribution of the overall phenotypicdiversity among sugarcane landraces in Ethiopia is unevenOur results showed that there was a wide range of variationthat existed in the local sugarcane landraces studied both atregional and within region levels Selection and differenti-ation of types might have taken place along a geographicalpattern perhaps associated with climate and use The presentstudy contributes to updating sugarcane descriptors adoptedfrom USDA-ARS as well as Bioversity passport data forthe future collection and evaluation As Ethiopian SugarCorporation is in the course of launching breeding programthe observed substantial variation of landraces would enablesugarcane breeders to design and practice breeding andselection programs to improve the crop
Data Availability
The data used to support the findings of this study areavailable from the corresponding author upon request
Conflicts of Interest
The authors declare that they have no conflicts of interest
Acknowledgments
The authors are grateful for the financial grant of SugarCorporation of Ethiopia Many thanks are due to EthiopianBiodiversity Institute (EBI) for its technical advice duringcollection of the local sugarcane genotypes Sincere thanksare forwarded to Ethiopian smallholder farmers who pre-served sugarcane germplasm and shared their knowledgeand experience Regional Zonal and District (ldquoWeredardquo)Agriculture Bureaus across Ethiopia are acknowledged fortheir support during the study
Supplementary Materials
These supplementary materials contain different tables sup-porting the results of the study These include passport dataof sugarcane germplasm collected and related informationon the materials local sugarcane germplasm collected andused formorphological diversity study introduced sugarcane
16 Advances in Agriculture
Table9Sh
anno
n-WeaverD
iversityIndices(H1015840)a
ndmeanandsta
ndarderror(plusmnS
E)of
land
racesineach
region
basedon
16discretecharacters
Characters
Amha
raBe
nsha
ngul-G
umz
Introd
uced
Oromia
SNNP
Tigray
ELb
Total
Presence
orabsenceo
fbud
cushion(BUDCUSH
ION)
083
095
095
089
082
098
090
088
Relativ
edegreeo
fbud
extension(BUDEX
TEND)
077
079
093
074
072
087
080
078
Relativ
ebud
Shape(BU
DSH
APE
)078
096
089
081
085
078
084
078
Canop
ystructure(C
ANOPY
)089
098
095
081
078
096
089
084
Relativ
eshape
ofdewlap(D
EWLA
PSHAP)
087
094
092
090
082
096
090
086
Relativ
eplant
erectness(ER
ECT)
080
082
050
067
071
088
073
064
Relativ
edegreeo
finterno
dealignm
ent(IN
ALIGN)
096
076
098
097
098
100
094
098
Relativ
einterno
deshape(IN
SHAPE
)086
069
095
077
083
091
083
082
Color
oftheleaves(LE
AFC
OLO
R)087
071
090
064
074
071
076
076
Type
ofou
tera
uricle(A
URI
CLE
OUT)
080
092
087
079
083
093
086
082
Relativ
eshape
ofligule(LIGSH
APE
)086
094
082
085
090
083
087
083
Presence
orabsenceo
fstalkcorkyc
racks(ST
ALK
CORK
C)
044
054
035
055
051
025
044
049
Presence
orabsenceo
fstalkcorkyp
atches(STA
LKCORK
P)066
000
072
079
059
087
060
069
Presence
orabsenceo
fstalkgrow
thcracks
(STA
LKCRA
CK)
066
100
088
062
088
099
084
080
Color
ofthee
xposed
rind(R
INDCOLE
)085
096
089
084
086
076
086
085
Presence
orabsenceo
fbud
groo
ve(BUDGRO
OVE)
095
100
072
100
100
099
094
100
Means
080plusmn0
03
081plusmn0
06
083plusmn0
04
079plusmn0
03
080plusmn0
03
085plusmn0
05
081plusmn0
03
080
b EL=
averageo
fentire
locatio
n
Advances in Agriculture 17
varieties used for morphological diversity study and fre-quency distribution for 16 qualitative characters in sugarcaneby regions in Ethiopia and country of sources of introducedgenotypes (Supplementary Materials)
References
[1] ldquoSpreading the word about clean solutions from sugarcanrdquoAccessed 04072018 httpwwwsugarcaneorg
[2] Central Statistics Agency (CSA) ldquoAgricultural sample surveyArea and production of major crops private peasant holdingsrdquoStatistical Bulletin Addis Ababa Ethiopia 2017
[3] T Esayas Exploration and Collection Characterization GeneticDiversity Analysis and Association of Traits for Yield and YieldComponents of Sugarcane (Saccharumspp) in Ethiopia [MasterThesis] Haramaya University Dire Dawa Ethiopia 2014
[4] J R Harlan Crops and Man American Society of AgronomyCrop Science Society of America Madison WI USA 2ndedition 1992
[5] R W Allard Principles of Plant Breeding JohnWilley and SonsNew York NY USA 2nd edition 1999
[6] B T Roach and J Daniels ldquoA review of the origin and improve-ment of sugarcanerdquo in Proceedings of Coper sugar InternationalSugarcane Workshop pp 1ndash32 1987
[7] P P Patil S S Patil and D U Patel ldquoGenetic diversity in sug-arcane (Saccharum spp Complex)rdquo Journal of Pharmacognosyand Phytochemistry vol 6 no 6 pp 570ndash573 2017
[8] R B Singh B Singh andR K Singh ldquoStudy of genetic diversityof sugarcane (Saccharum) species and commercial varietiesthrough TRAP molecular markersrdquo Indian Journal of PlantPhysiology vol 22 no 3 pp 332ndash338 2017
[9] R W Allard ldquoPopulation structure and sampling methodsrdquo inGenetic Resources in PlantsmdashTheir Exploration and Conserva-tion O H Frankel and E Bennett Eds IBP Handbook no 11pp 97ndash107 FA Davis Company Philadelphia PA USA 1970
[10] D R Marshall and A H D Brown ldquoOptimum samplingstrategies in genetic conservationrdquo inCrop Genetic Resources forToday and Tomorrow O H Frankel and J G Hawkes Eds pp53ndash80 Cambridge University Press 1975
[11] J G Hawkes Crop Genetic Resources Field Collection ManualPudoc Wagnningen Netherland 1980
[12] J G Hawkes The Diversity of Crop Plants Harvard UniversityPress London England 1983
[13] B P K Yerima Manual for Good Laboratory Practice Foodand Agriculture Organization of United Nation Addis AbabaEthiopia 1993
[14] A Walkley and I A Black ldquoAn examination of the degtjareffmethod for determining soil organic matter and a proposedmodification of the chromic acid titrationmethodrdquo Soil Sciencevol 37 no 1 pp 29ndash38 1934
[15] S R Olsen and L E Sommers ldquoPhosphorusrdquo in AgronomyMonograph A L Page et al Ed Methods of soil analysis part2 pp 403ndash430 ASA and SSSAMadisonWI USA 2nd edition1982
[16] R H Bray and L T Kurtz ldquoDetermination of total organic andavailable forms of phosphorus in soilsrdquo Soil Science vol 59 no1 pp 39ndash46 1945
[17] S Sahlemedhin andB Taye ldquoProcedures for soil and plant anal-ysisrdquo Tech Rep 74 National Soil Research Center EthiopianAgricultural Research Organization 2000
[18] F C Stickler S Weaden and AW Pauli ldquoLeaf area determina-tion in grain sorghumrdquo Journal of Agronomy and Crop Sciencevol 53 pp 187-188 1961
[19] Munsell Color ChartThe Munsell Book of Color Matte Collec-tion Munsell Color Chart NewWindsor NY USA 1990
[20] H O Hartley ldquoThe maximum F-ratio as a short cut test forheterogeneity of variancesrdquo Biometrika vol 37 pp 308ndash3121950
[21] L Pecetti P Annicchiarico and A B Damania ldquoBiodiversity ina germplasm collection of durumwheatrdquo Euphytica vol 60 no3 pp 229ndash238 1992
[22] L Pecetti and A B Damania ldquoGeographic variation intetraploid wheat (Triticum turgidum ssp turgidum convardurum) landraces from two provinces in Ethiopiardquo GeneticResources and Crop Evolution vol 43 no 5 pp 395ndash407 1996
[23] S K Jain C O Qualset G M Bhatt and K K Wu ldquoGeo-graphical patterns of phenotypic diversity in a world collectionof durumwheatsrdquoCrop Science vol 15 no 5 pp 700ndash704 1975
[24] K K Nkongolo L Chinthu M Malusi and Z Vokhiwa ldquoPar-ticipatory variety selection and characterization of sorghum(Sorghum bicolour (L) Moench) elite accessions from Malaw-ian gene pool using farmer and breeder knowledgerdquo AfricanJournal of Agricultural Research vol 3 pp 273ndash283 2008
[25] M B Semira ldquoInnovative methods and metrics for agricul-ture and nutrition actions soil quality and human nutritionin Ethiopiardquo Leverhulme Center for Integrative Research onAgriculture and Health (LCIRAH) 2017
[26] I A Khan A Khatri M A Siddiqui G S Nizamani and SRaza ldquoPerformance of promising sugarcane clone for yield andquality traits in different ecological zones of Sindhirdquo PakistanJournal of Botany vol 36 no 1 pp 83ndash92 2004
[27] M F Perera M E Arias D Costilla et al ldquoGenetic diversityassessment and genotype identification in sugarcane based onDNAmarkers andmorphological traitsrdquo Euphytica vol 185 no3 pp 491ndash510 2012
[28] C G Neil M Kay and C C Jack ldquoDiversity among mainlandUSA sugarcane cultivars examined by SSR genotypingrdquo Journalof the American Society of Sugar Cane Technologists vol 29 pp36ndash52 2009
[29] S SchenckMWCrepeauK KWu P HMooreQ Yu andRMing ldquoGenetic diversity and relationships in native HawaiianSaccharum officinarum sugarcanerdquo Journal of Heredity vol 95no 4 pp 327ndash331 2004
[30] R Gebretsadik H Shimelis M D Laing P Tongoona andN Mandefro ldquoA diagnostic appraisal of the sorghum farmingsystem and breeding priorities in Striga infested agro-ecologiesof Ethiopiardquo Agricultural Systems vol 123 pp 54ndash61 2014
[31] L E V Cesar A T Bruzi J A R Nunes et al ldquoGeneticdivergence of sugarcane varieties collected in the region ofLavras Minas Gerais Brazilrdquo Genetics and Molecular Researchvol 14 no 4 pp 14043ndash14049 2015
[32] C Grenier P J Bramel J A Dahlberg et al ldquoSorghums of theSudan analysis of regional diversity and distributionrdquo GeneticResources and Crop Evolution vol 51 no 5 pp 489ndash500 2004
[33] N V Nair R Balakrishnan and T V Sreenivasan ldquoVariabilityfor quantitative traits in exotic hybrid germplasm of sugarcanerdquoGenetic Resources and Crop Evolution vol 45 no 5 pp 459ndash464 1998
[34] R Nagarajan S Alarmelu and R M Shanthi ldquoStudies onvariation in interspecific hybrids of Saccharumrdquo Sugar Tech vol2 no 3 pp 42ndash46 2000
18 Advances in Agriculture
[35] D N Kamat and J R P Singh ldquoVariability in Sugarcane underrain fed conditionrdquo Sugar Technology Short Communicationsvol 3 no 1-2 pp 65ndash67 2001
[36] B S Rishipal A S M Chaudhary and S P Kadian ldquoStudieson character interrelationship in clonal generation of sugarcane(Saccharum complex hybrids)rdquo Indian Sugar vol 47 no 11 pp907ndash911 1998
[37] S Thippeswamy S T Kajjidoni P M Salimath and J V GoudldquoCorrelation and path analysis for cane yield juice quality andtheir component traits in sugarcanerdquo Short Communication vol5 no 1-2 pp 65ndash72 2003
[38] P S Verma S Pal and N K Karma ldquoGenetic variability andcorrelation studies in sugarcanerdquo Indian Sugar vol 49 pp 125ndash128 1999
[39] H A Esechie J W Maranville and W M Ross ldquoRelationshipof stalk morphology and chemical composition to lodgingresistance in sorghumrdquo Crop Science vol 17 no 4 p 609 1977
[40] P K Das B C Jena and N Nayak ldquoCharacter association andpath analysis of sugar yield in sugarcanerdquo Indian Sugar vol 46no 10 pp 805ndash808 1997
[41] K J Sreekumar M Kuriakose M Thomas D Alexander andS Santhakumari ldquoVariability heritability and correlation onstudies on the yield and quality characters of sugarcanerdquo IndianSugar vol 44 no 4 pp 243ndash249 1994
[42] J R Harlan J M de Wet and E G Price ldquoComparativeevolution of cerealsrdquo Evolution vol 27 no 2 pp 311ndash325 1973
[43] A Elings ldquoDurum wheat landraces from syria II patterns ofvariationrdquo Euphytica vol 54 no 3 pp 231ndash243 1991
[44] K Tato ldquoRainfall in Ethiopiardquo Ethiopian Geographical Journalvol 2 pp 28ndash36 1964
[45] A B L Stemler J R Harlan and J M J deWet ldquoThe sorghumsof ethiopiardquo Economic Botany vol 31 no 4 pp 446ndash460 1977
[46] P Annicchiarico L Pecetti and A B Damania ldquoRelationshipsbetween phenotypic variation and climatic factors at collectingsites in durum wheat landracesrdquo Hereditas vol 122 no 2 pp163ndash167 1995
[47] A B Damania L Pecetti C O Qualset and B O HumeidldquoDiversity and geographic distribution of adaptive traits inTriticum turgidum L (durum group) wheat landraces fromTurkeyrdquoGenetic Resources and Crop Evolution vol 43 no 5 pp409ndash422 1996
[48] A Ayana and E Bekele ldquoGeographical patterns of morpho-logical variation in sorghum (Sorghum bicolor (L) Moench)germplasm from Ethiopia and Eritrea quantitative charactersrdquoEuphytica vol 115 no 2 pp 91ndash104 2000
[49] D S Falconer Introduction to Quantitative Genetics JohnWilley and Sons Inc New York NY USA 3rd edition 1993
[50] R S Thorpe ldquoBiochemical analysis of geographical variationand racial affinitiesrdquo Biology Review vol 51 pp 407ndash452 1976
[51] M Farooq ldquoMorphological and agricultural characteristicsof sugarcane in relation to mechanized agriculture at NARCIslamabadrdquo Portu Rico Journal vol 1 pp 51ndash61 1989
[52] M Akhtar N N Elahi and M Ashraf ldquoEvaluation of exoticsugarcane varieties for agronomic characters and productivityrdquoPakistan Journal of Biological Sciences vol 4 no 1 pp 37ndash402001
[53] F R Piscitelli ldquoPrincipals caracteres exomorfologicos de cul-tures de cana de azucarrdquo TUC Iindustraly Agricos vol 7 pp 49ndash57 1994 (Spanish)
[54] A H D Brown and B S Weir ldquoMeasuring genetic variability inplant populationsrdquo in Isozyme in Plant Genetics Breeding S DTanksley and T J Orton Eds part A Elsevier Science 1983
[55] R Balakrishnan N V Nair and T V Sreenivasan ldquoA methodfor establishing a core collection of Saccharum officinarum Lgermplasm based on quantitative-morphological datardquo GeneticResources and Crop Evolution vol 47 no 1 pp 1ndash9 2000
[56] C A Nestor and B Q Reynaldo Sugarcane Variety Improve-ment in Southeast Asia and the Pacific for Enhanced andSustainable Productivity ndash Germplasm and Disease IndexingComponents Institute of Plant Breeding (IPB) College ofAgriculture (CA) UP Los Banos (UPLB) 2003
[57] A Maqbool J Muhammad and A Sagheer ldquoAgronomic traitsand morphological characteristics of some exotic varieties ofsugarcanerdquoPakistan Journal of Agricultural Research vol 19 no4 2006
by the leading sugar producer Brazil with a total amount of353 million tons produced in 201617 [1] Similarly Brazilianethanol production reached 3023 billion litres in 201516
Though commercial sugarcane production has a historyof six decades sugarcane had been cultivated in Ethiopiasince 16th century According to the report by central statisticsagency (CSA) currently sugarcane is produced in about3123681ha with 156506000 holdings in different parts ofthe country [2] But the production is not usually usedfor industrial purposes It is noticeably used for makingconfectioneries household consumption (chewing) sellingfor immediate cash and feeding livestock In some areassugarcane is used to prepare local beverage called ldquoKaribordquomainly preferred byMuslim communities while in others theleaves are used for thatching and as firewood [3] Howeverthe potential of this sector is not well explored and has notbeen given due consideration Furthermore no explorationand germplasm collection have been done to represent andpreserve local landraces
The sugar industry of Ethiopia is so far dependent onintroduction of exotic varietieswhich are not suitably adaptedto various agroecologies and local growing conditions Inlight of the rapidly increasing commercial sugarcane planta-tion areas in the country the demand for improved varietiesthat suit various agroecologies is increasing Under suchsituations there will be a continuous demand for broadgenetic base sugarcane varieties that are high yielding andstable under abiotic and biotic stressesTherefore the industryis currently launching breeding program which is long over-due to produce its own improved varieties The developmentof high yielding and stable varieties requires a continuoussupply of new germplasm as a source of desirable genesandor gene complexes The primary sources of such genesare landraces introductions weedy and wild relatives of cropplants [4 5] The availability of such germplasm requiresthe identification of areas of diversity of various charactersof agronomic importance especially in the local landracesgrowing within the variable agroecologies of Ethiopia There-fore germplasm collection and conservation and the studyof genetic diversity of Ethiopian sugarcane landraces areworthwhile since this can broaden the genetic base andprovide locally adapted genes for improvement of the crop
In spite of the great importance of Ethiopian sugarcanelandraces for the germplasm genetic base improvement andutilization in the breeding program no effort has beenmade so far to collect and preserve this genetic wealthFurthermore study of the variation and assessment of extentand geographical pattern of distribution of this landracesis lacking For effective utilization of germplasm in plantbreeding programmes the information on the extent andpatterns of distribution of genetic variation of a crop species isvery essential [6ndash8]This also canhelp in devising appropriatesampling procedures for germplasm collection and conser-vation and obtaining core collection for efficient germplasmmanagement [9 10]
The objectives of this study were to collect sugarcanelandraces in Ethiopia and to assess phenotypic diversity andanalyze regional distribution among landraces collected fromdifferent geographical areas
2 Materials and Methods
21 Germplasm Collection
211 Sampling Technique Sugarcane germplasm were col-lected during 201011 all across Ethiopia in the regionalstates of Amhara (07102010-09052011) Oromia (26082010-08072011) Southern Nations Nationalities andPeoples Region (SNNPR) (10082010-09122010) Tigray(21042011-09052011) Benshangul-Gumz (20122010-15022011) Gambella (04112010-28112010) Somali (29062011-26072011) and Harari (02062011-23062011)(Figure 1 Supplemental Table 1) Collection was made fromhomesteads farmersrsquo fields and local markets Germplasmwas collected using stratified random sampling techniquesampling areas are shown in Figure 1 In each region all zoneswere sampled Two to four districts (locally referred to asldquoWeredasrdquo) were selected from each zone From each district(depending on size a district contains several localities orsubdistricts) 2-5 subdistricts (locally referred as ldquoKebelesrdquo)or peasant associations (PAs) were selected The districts andsubdistricts were selected based on long agricultural historyand relatively wide areas allocated to sugarcane productionMoreover purposive sampling was also employed based oninformation supplied by key informants on the unique andquality sugarcane types grown in these areas In the selectedsubdistricts sugarcane clones were collected following themethods proposed in [11] Each distinct morphotype in avillage was randomly sampled Information on the sampledsugarcane germplasm was recorded and passport data wascollected following the method of Bioversity International[12] Moreover juice sample was taken from the bottommiddle and top part of the stalk of each clone and meanpercent brix reading was recorded using hand refrac-tometer
212Determination of the Physical andChemical Properties ofSoils in SugarcaneGermplasmCollectionAreas Todeterminethe predominant physical and chemical properties and thefertility status of soils under sugarcane production sampleswere collected and analyzed across germplasm collectionareas Georeferencing (latitude and longitude) of the studysites were made with a Garmin GPS In every germplasmsampling area a composite soil sample was taken between0ndash30 and 30ndash60 cm depths Soil samples were analyzed fororganic carbon total nitrogen (N) soil pH soil electricalconductivity (EC) available phosphorus (P) and availablepotassium (K) contents using standard procedures [13] SoilpH was measured potentiometrically using a digital pHmeter (Jenway Model-3320 GransmoreGeeen) Soil EC wasmeasured using digital conductivity meter (Jenway Model-4310 GransmoreGeeen) Organic carbon was determinedfollowing the wet digestion method described in [14] Total Nwas determined using the Kjeldahal procedure [13] Availablephosphorus was determined using the Olsen method [15]and Bray II method [16] for acidic soils Available K wasmeasured by flame photometry using the sodium acetateextractant method at pH 48 [17] Soil texture was determinedby hydrometer method [17]
Advances in Agriculture 3
1700000
1500000
1300000
1100000
900000
700000
500000
300000
1700000
1500000
1300000
1100000
900000
700000
500000
300000
minus300000 100000 500000 900000 1300000 1700000
minus300000 100000 500000 900000 1300000 1700000
Sampling Site Location Map
Figure 1 A map of Ethiopia showing sugarcane germplasm collection sites (black circles)
Table 1 Location information of the test sites
Location Latitude Longitude Altitude (masl)119886 Rainfall (mm) Temperature (∘C)Minimum Maximum
Wonji 8∘311015840 N 39∘121015840 E 1550 800 153∘C 269∘CMetehara 8∘511015840 N 39∘ 521015840 E 950 554 175∘C 326∘C119886masl = meters above sea levelSource meteorological Stations of respective locations
22 Diversity Study
221 Plant Materials A total of 211 sugarcane (Saccharumspp) accessions consisting of 196 landraces (SupplementalTable 3) and 15 introduced commercial varieties including2 standard varieties (Supplemental Table 4) were used forthis study The landraces represent collections from allgeographical regions across Ethiopia maintained at fieldconservation garden at Wonji and Metehara Sugar EstatesSugar Corporation of Ethiopia Sampling of the landraces wasmade on representation basis stratified systematic samplingmethod to a given range of geographic area altitudinalranges and a range of morphological traits The introducedmaterials were commercial varieties under production indifferent estates where the two were standard varieties
The 196 landraces were collections from the followingregions of Ethiopia Amhara (47) Benshangul-Gumz (10)Gambella (3) Harari (2) Oromiya (65) SNNPR (59) Somali(3) and Tigray (7) (Figure 1) The altitude of the collectionsites for the landraces used in this study ranged from 454to 2687 meters above sea level representing the distributionof the crop in Ethiopia The introduced materials were fromBarbados (6) Cuba (1) India (5) Mexico (1) and SouthAfrica (2)
222Methods The plant materials were grown at Wonji andMetehara Sugar Estates of Sugar Corporation of Ethiopiaduring the 201213 growing season Details of the plantinglocations are shown in Table 1 Each accession was grown ina single row plot of 5m long and 145m between rows and20 cm between plants within a row with two replications inrandomized complete block design
Uniform crop management practices were applied asrecommended for the areas Urea was applied 25 monthsafter planting at a rate of 200 kgsdothaminus1 atWonji and 400 kgsdothaminus1at Metehara
Accessions from regions with sample size less than 12were included in adjacent regions to reduce experimentalerror due to small sample size Hence the two Harari andthree Somali accessions were included in Tigray and thethree Gambella accessions were included in Benshangul-GumzThis reduced the 8 regions of Ethiopia fromwhich thelandraces were originally drawn to five With the introducedmaterials included 6 regions of origin were used in thestatistical analyses
Data on 17 quantitative stalk characters was recordednamely sprout count 1 and 2 months after planting(SPC1MAP and SPC2MAP) tiller counts 4 and 5month afterplanting (TC4MAP and TC5MAP) stalk count 10 months
4 Advances in Agriculture
after planting (STC10MAP) hand refractometer brix reading10 months after planting (HRBrix10MAP) millable stalkcount per hectare (MSCHA) single cane weight (SCW)number of internode (NOI) internode length (IL) stalkheight (SH) stalk diameter (SD) leaf length (LL) leaf width(LW) leaf area (LA) cane yield quintal per hectare (CYHA)and sugar yield quintal per hectare (SY) Data on 4 juicequality parameters ie brix percent (Brix) pol percent(pol ) purity percent (purity ) and sugar percent (SR) was also recorded For every accession ten plants wereused for recording data for quantitative characters whichwere recorded on plot basis Count data and cane yield wererecorded considering all cane stalks from the whole plot Forquantitative leaf characteristics measurement a proceduredeveloped in [18] was used
To categorize each accession morphologically sugarcanedescriptors adopted fromUSDA-ARSwere employed (GRIN2004) Data on 16 qualitative traits was recorded namelypresence or absence of bud cushion (BUDCUSHION) rel-ative degree of bud extension (BUDEXTEND) relative budshape (BUDSHAPE) relative shape of dewlap (DEWLAP-SHAP) type of outer auricle (AURICLEOUT) presence orabsence of stalk corky cracks (STALKCORKC) presenceor absence of stalk corky patches (STALKCORKP) relativeshape of ligule (LIGSHAPE) presence or absence of stalkgrowth cracks (STALKCRACK) presence or absence of budgroove (BUDGROOVE) relative plant erectness (ERECT)relative degree of internode alignment (INALIGN) relativeinternode shape (INSHAPE) colour of the leaves (LEAF-COLOR) colour of the exposed rind (RINDCOLE) andcanopy structure (CANOPY) Each accession was scored forthe most frequent character-state Leaf colour and colourof the exposed rind were examined and scored using theMunsell colour chart [19]
223 Statistical Analyses First analysis of variance wasmadefor the 21 characters for each location Homogeneity ofthe error variances among the locations was assessed in[20] F-max method for each of the 21 characters The testestablished the homogeneity of the error variances for allcharacters except TC4MAP For TC4MAP logarithmic datatransformation which is recommendable for continuousdata was used to homogenize the error variance Thendata of all the characters were subjected to test location-accession-related analysis of variance to determine effects oftest sites and genotype X environment (G X E) interactionThe results showed the nonsignificance of effects of testsites for SPC1MAP IL LW Pol sugar percent (SR) andthe nonsignificance of G X E interactions for SPC2MAPSCW Brix Pol and Brix10MAP For the other charactersboth test site effects and G X E interactions were significant(Plt005)
Analysis of variance was made for 21 quantitative char-acters following the procedure used in [21 22] The meansquares of the regions were tested against pooled meansquares of accessions within regions The pooled meanssquares for accessions within regions of origin and the meansquares of accessions within each region were tested against
the pooled within-region error mean squares Means rangesfor means and percent coefficients of variation for all thecharacters were computed for each region of origin and forthe entire data The regional means were compared usingDuncanrsquos multiple range testing Correlations between thecharacters were computed at three levels First correlationsof the characters were assessed based on the 211 accessionmeans Then interregion correlation was computed usingthe means of characters for each region Finally a series ofintraregion correlation coefficient matrices were obtained foreach region using the accession means from that region forthe characters
For qualitative characters phenotypic frequency distri-butions were worked out for all the sample germplasm andlocations The ShannonndashWeaver diversity index (H1015840) wascomputed using the phenotypic frequencies to assess thephenotypic diversity for each character for all accessions TheShannonndashWeaver diversity index as described in [23] is givenas
1198671015840 = minus119899
sum119894=1
119901119894log119890(119901119894) (1)
where pi is the proportion of accessions in the 119894th class of ann-class character and n is the number of phenotypic classesof traits Each 1198671015840 value was divided by its maximum value(logen) and normalized in order to keep the values between0 and 1 By pooling various characters across the regions theadditive properties of 1198671015840 were used to evaluate diversity ofregions and characters within the population The averagediversity index (H1015840) over n traits was estimated as H1015840 =sumH1015840n
3 Results and Discussion
31 Collection of Sugarcane Germplasm In Ethiopia thehistory of sugarcane cultivation by smallholder farmers pre-ceded that of commercial cultivation As documented in thehistory of a monastery in Northern Ethiopia sugarcane hasbeen grown in the country since the 16th century [2] In thisstudy local sugarcane germplasm exploration and collectionwere conducted all across Ethiopia in the regional statesMore than 300 local sugarcane genotypes were collectedduring 20102011 and passport data of the genotypes ispresented in Supplemental Table 1 The collected germplasmwere planted at five locations across the country with respectto their collection area These are Wondogenet AgriculturalResearchCenter JimmaAgricultural ResearchCenterMechaWereda Agricultural Bureau nursery field at Picolo AbaySirinka Agricultural Research Center (at Kobo Subcenter)and research field of Haramaya University at Diredawa(Tonny Farm) The clones have been monitored during the201112 season for symptoms of major diseases and insectsThese clones were transferred to commercial sugarcane plan-tation estates at Wonji and Metehara for further selectionand maintenance No major diseases and insect pests weredetected in all the germplasm collected except sometimestermites and borer were observed (Supplemental Table 2)
Advances in Agriculture 5
Table 2 Mean soil physical and chemical properties of sugarcane fields in the surveyed regions
Regions Soil chemical properties Texture ()PH EC(dsm) OC N P (ppm) K(ppm) Sand Silt Clay
Most of the sampled germplasm had acceptable levels of juicerefractometer readings (expressed in degree brix) relative tothe history of the ages of the samples (Supplemental Table 1)The number of harvestablesaleable cane stalks obtained inplant cane and ratoon crops and the number of harvestableratoon crops showed variation region to region and zone tozone and sometimes district (ldquoweredardquo)-wise There is alsovariation in maturity time of the clones both in plant caneand in ratoon crops (Supplemental Table 2)These are mainlydepending on the variety soil condition cultural practicesand climatic conditions
The genotypes collected from the respective zones inthe study regions were recorded by their local namesSometimes similar varieties bear different names at differentlocalities and vice versa Farmers mentioned a broad rangeof sugarcane landraces that had been grown in the areasand maintained for generations Some of the landraces werecommonly recognized by most farmers within and acrosszones of the respective regions whereas some were rarevarieties known only by few farmers The large number oflandraces observed during the current study demonstratesthe existence of diverse genetic resources of sugarcane inEthiopia The diversity could have evolved presumably due todiverse climatic conditions low input management systemshigh pests and disease pressure and continuous selectionby farmers Smallholder farmers face diverse environmentstresses and have multiple production objectives that affectselection of genotypes [24] There were no formally releasedimproved sugarcane varieties grown by farmers in the studyregions except in few places This was mainly observedin SNNPR at Wolayta and Kembata-Tembaro zones Thesevarieties have been informally introduced to these areas byseasonal workers employed at the sugar estates especially atWonji and Metehara
The collected landraces would serve to broaden thegenetic base of the available sugarcane germplasm Thesugar industry in Ethiopia is currently establishing sugarcanebreeding programTherefore collection and efficient charac-terization of germplasm are vital to strengthen the breedingprogram
311 Physical and Chemical Properties of Soils of Sugar-cane Germplasm Collection Areas Table 2 summarizes thephysical and chemical properties of soils in the sugarcane
production fields of surveyed areas Sugarcane productionby smallholder farmers in the surveyed areas is conductedacross the seasons where harvesting and planting are done aslong as there is available moisture Therefore the soil samplesrepresent the sugarcane production season and the altitudinalranges of the respective areas
Generally the organic carbon content of soils in sugarcaneproducing areas across the country is low to very low (2-4lt2) Amhara Gambella SNNP Somali and Tigray Regionsare in a very low category (lt2) Similarly across regions thetotal N content is low except Harari and Oromia regionswhere the average total N is medium This problem shouldbe addressed either with augmenting the soil with inorganicfertilizers or through appropriate management practices likecrop residue management Available phosphorus is high inAmhara Gambella and SNNP Regions whereas moderatevalues were observed in Harari Oromia and Tigray RegionsGenerally the level of phosphorus in soils of sugarcaneproducing areas in these regions is adequate In Benshangul-Gumz and Somali Regions low level of phosphorus wasrecorded which should be addressed with due attentionAvailable K is sufficient for normal sugarcane growth in soilsof all regions Relatively lower values of K were recorded insoils of Harari and Tigray Regions It should be noted thatorganic carbon EC PH and variability of P andK are affectedby rainfall and temperature that are seasonal and the valuesrecorded refer to the seasonwhen the soil samples were takenGenerally Ethiopian soils are deficient in various essentialnutrients like boron nitrogen phosphorus potassium sulfurzinc and copper although severity differs from region toregion [25] Particularly the loss of P and N resulting fromthe use of dung and crop residues for fuel is demonstrated tobe equivalent to the total amount of commercial fertilizer use
Across regions in Ethiopia generally clay type of soildominates followed by silt except in Gambella Region wherethe dominant soil texture was sand Soil management optionsrelated to clay soil may address soil related problems insmallholder sugarcane growing areas across Ethiopia
32 Diversity of Sugarcane Landraces
321 Quantitative Characters
Univariate Statistics Analysis of variance depicted highlysignificant differences (p lt 001) between accessions pooled
6 Advances in Agriculture
over the regions for the 21 characters of the 211 sugarcaneaccessions studied and significant difference for the 6 regionsof origin for 6 characters (Table 3) The results suggestedthe occurrence of significant phenotypic variation betweenthe accessions as a whole Significant variations of differentsugarcane stalk and juice quality characters was also reportedin similar studies conducted elsewhere [26 27] Region-wise partitioning of the variance indicated significant within-region differences (p lt 005) among the populations withinalmost all regions for the characters NOI IL SH SD LLLW LA Brix Pol Purity SR MSCHA SCW CYHAand SY for 8 characters within Tigray for TC4MAP andTC5MAP within Oromiya and SNNPR Brix10MAP showedno significant difference within all regions
In general within-region variation was greater for stalkdiameter single cane weight millable stalk count and caneand sugar yield than for other characters for all the regionsAssuming that a significant portion of the phenotypic vari-ation is genetic it would be possible to make selection forany of this group of characters within a particular regionIt was apparent that the variance between accessions pooledover regions was greater than between regions Therefore inorder to sample the variation effectively it would be necessaryto sample the variable populations from different localitiesin a region The study in [28] on genetic diversity amongmain land USA sugarcane cultivars showed high geneticdiversitywithin populations of different regions than betweenpopulations
Duncanrsquos multiple range testing for regional means for allthe characters is shown in Table 4 Relatively much differenti-ationwas observed for stemdiameter and juice purity percentcompared to other parameters Important yield componentslike MSCHA SCW SH and SD have shown higher valuesin Tigray SNNPR and Oromiya regions High cane andsugar yield were recorded for accessions in Tigray regionfollowed by SNNPR Oromiya and introduction which werestatistically at par Statistically similar lower values of caneand sugar yield were recorded in Amhara and Benshangulcollections
Higher value of sucrose percent in cane was also observedfor accessions from Tigray and the lowest value was inAmhara accessions while other regions were statisticallysimilar Generally for most of the characters high valuewas obtained from accessions in Tigray region followed bySNNPROromiya and introduction Therefore the materialsfrom these regions can be used for selection of cane and sugaryield per se and in breeding program The local accessions inmany of the regions showed superiority over the introducedcommercial varieties currently in production for most of thecharacters including cane and sugar yield (Table 4)This maybe due to the adaptation and coevolution of the materialswith different stresses for long period of time in the countryrsquosagroecology These locally adapted genes conferring yieldadvantage could be harnessed through strategic crossing andselection for improvement of the crop Superiority of locallandraces over introduced varieties was reported [29 30]
The range of variation of the accession means establishedwide variation between the regions and the accessions withinthe regions for the characters studied (Table 5) Accordingly
the maximum score was 41 times the minimum for sproutcount one month after planting 32 for sprout count twomonths after planting 25 for tiller count four months afterplanting 22 for sugar yield 20 for cane yield 12 for millablestalk count 8 for stalk count 10 months after planting 7for single cane weight 6 for tiller count five months afterplanting and 2 for other yield components and sugar qual-ity parameters except hand refractometer brix reading tenmonths after planting and juice purity percent which scored1 The same trend was observed between accessions withina particular region However accessions within AmharaBenshangul-Gumz and introduction showed wider ranges ofvariation than those accessions within other regions for themajority of the characters The wide range of geographic andclimatic features of the regions in Ethiopia and the originaldifferentiation and further farmers selection for productionniches and uses for hundreds of years may have resultedin the accessions to possess a tremendously high degree ofmorphological variation The reason for introduced varietiesshowing high variation also could be due to broader geo-graphic spectrum fromwhere theywere initially acquired anddifference in their parental source and genealogical historyIt was demonstrated even with these limited numbers ofsamples (15 genotypes) that introduction of genetic materialshas practically broadened the genetic base of the sugarcanegermplasm for the breeding program
Coefficient of variation measured as the ratio of standarddeviation to the corresponding overall mean and expressedas percentage is useful to compare different charactersmeasured in different units It is also useful to compare samecharacter in different groups of populations with differentsample size mean and variance or different characters indifferent populations In the current study high coefficientsof variation were observed between regions and within eachregion for sprout as well as tiller counts in the monthsthe data was recorded stalk count 10 months after plant-ing and millable stalk count at harvest cane and sugaryield single cane weight and leaf area (Table 6) Thoughaccessions from a particular region were more variable fora specific character compared to other regions accessionsfromAmhara Benshangul-Gumz introduction and SNNPRwere more variable than accessions from other regions Thisresult demonstrated the tremendous variability of sugar-cane germplasm from these regions The accessions fromTigray and Oromiya had low coefficients of variation formany characters indicating relatively high within-regionuniformity The high coefficients of variation observed formost of the characters agreed well with those reportedin [31] for indigenous sugarcane of Brazil Similar resultswere also reported in sorghum [32] The different levels ofregional variability of a particular character could be due todifferences in forces of selection andor differences in theintensity of a particular selecting force It could also be dueto random effect and reduced sample parents of parentalmaterials moved differently by various human populationmigration and the interaction of these with Darwinianforces
Results of this study suggested the presence of wide rangeof variations of characters determined by univariate statistics
This is in agreement with previous studies on sugarcanegermplasm [33ndash35]
This is the first study to report information on geo-graphical pattern of variation in Ethiopian local sugarcanegermplasm which has been lacking so far Our results pro-vided experimental evidence on occurrence of geographicalvariation and significant within-region variation where it washigh in the regions such as Amhara Benshangul-Gumz andSNNPR
The overall patterns of similarity or difference betweenregions seemed to depend on environmental factors such asrainfall temperature length of growing season and altitudeWide variability of millable stalk count at harvest singlecane weight and plant height was observed among regionswhere these characters demonstrated high correlation withcane and sugar yield (Table 7) Therefore breeders can utilizeaccessions of regions showing variability for these charactersin selection programs and to design breeding strategies toproduce varieties with best commercial merits
Bivariate Statistics Correlation coefficients worked out onthe 21 quantitative characters are shown for the entire data(Table 7) between regions (Table 8) and within regions(data not shown) Results of the association studies for theentire data showed that cane and sugar yield had highlysignificant positive correlation with most of the quantitativestalk characters and juice quality parameters However theassociation of tiller counts 4 and 5 months after plantingstalk count 10 months after planting millable stalk countat harvest single cane weight and plant height with caneand sugar yield was stronger suggesting these componentcharacters asmain contributing factor to cane and sugar yieldIn fact sugar yield is the product of cane yield and sucrose per-cent cane surely being influenced with components affectingcane yield Similar trend was reported by earlier sugarcaneworkers [36 37] In sugarcane the cane and sugar yields areconsidered to be the complex characters The information onthe phenotypic interrelationship of cane yield and sugar yieldwith their component character per se would be of immensehelp to sugarcane breeders
Stem diameter had significant positive association withboth cane and sugar yield indicating the significance ofthis trait in improving both cane and sugar yields Similarreports have beenmade by earlier sugarcane workers [37 38]Stalk diameter was also positively and significantly correlatedwith plant height It is one of the traits that are related tolodging resistance [39] and the positive associations withheight would help in reducing the chance of lodging as heightincreases Among the physiological attributes included in thestudy leaf area had highly significant positive associationwith cane yield Similar observation was also made in [37]Sucrose percent which is an important juice quality attributehad highly significant positive association with sugar yieldand simultaneously cane yield also had highly significantassociation with sugar yield Based on the magnitude ofcorrelation coefficient values it can be said that cane yield ismuch more important than sucrose percent in determiningthe sugar yield and the same has also been reported by othersugarcane workers [37 40] Similar to observations made in
[37 41] results of the current study showed that all the qualityparameters like sucrose percent pol percent brix percentand purity percent were significantly correlated in positivedirection This indicated any of these juice quality traitscould be considered for selection leading to the simultaneousimprovement in the remaining quality traits
Ten of the 21 characters also showed significant positivecorrelations with altitude of the collection sites (Table 7) Asdescribed in [42] ecological characteristics have influencedthe genotypic constitution of landraces during domesticationand hence a relationship exists between the agroecology inthe collection site and the morphological characteristics ofthe landraces Thus positive correlation between collectionsite variables and plant characteristics would imply that thevariation between accessionsmay be related to agroecologicalvariations among the collection sites [43]
The correlation coefficient between number of internodesand altitude was negative and significant indicating thatother environmental factors (other than altitude) andornonenvironmental factorsmight account for the variation forthis particular character Since temperature decreases withan increase of altitude in Ethiopia [44 45] it is more likelythat temperature has exerted strong selection pressure innumber of internodes in the genotypes Information on therelationships between environmental factors of the collect-ing sites and morphophysiological variation of germplasmcould enhance the understanding of evolutionary adaptivepatterns which could assist germplasm collectors and users[46]
Character associations between region and within regionalso followed similar fashion like that of the entire dataNumber of sprout tiller and stalk counts and sugar qualityparameters showed significant positive correlations with caneand sugar yield
Though not significant and strong the correlation of leaflength and width and leaf area (leaf area = leaf length Xleaf width X 075) with cane and sugar yield was negativein the interregion correlations This kind of relationship wasalso exhibited between these leaf characteristics and yieldcomponents like tiller and stalks counts plant height andsingle cane weight and stalk diameter and sugar qualityparameters This may have been attributed to the variationof temperature with altitude and other environmental factorsacross the regions resulting in such type of relationshipSuch relationship also calls for the need to have enough dataon environmental variables of the collection sites such astemperature and rainfall if germplasm collection is requiredtomeet its objectives effectively and efficiently Unfortunatelydata on such useful environmental variables like temperatureand rainfall are lacking in the passport data of the currentEthiopian sugarcane germplasm collections The significanceof environmental factors of the collection sites as veryimportant determinants in structuring morphophysiolog-ical variations was reported in tetraploid wheat [22 4748]
Association of characters among yield its componentsand other economic traits is important for the interpretationof the patterns of variation and making selection in breed-ing program and combining several desirable attributes It
suggests the advantage of a scheme of selection for morethan one character at a time The correlation between char-acters may arise from linkage or from developmental geneticinteractions with orwithout a purely phenotypic componentIt could arise also due to differential phenotypic plasticityof characters themselves Within the limit of experimentalerror and environmental effects high correlation coefficientsbetween characters may show that the characters sharesome common element of genetic control (ie pleiotropylinkage) between genes or else from independently controlledcharacters responding similarly to geographic variation inselection pressures [49] The interregion correlation coef-ficient between the characters measures the consistency oftheir patterns of regional variation while the intraregioncorrelation coefficient measures the association arising fromgenetic factors but is not affected by regional variation[50] Since this study showed significant positive correlationsintraregionally for some character combinations it wouldseem that common genetic control might be playing a role inbringing about correlations between the various charactersIt appeared that different response to regional variationwas playing a greater role than different genetic controlas shown by the many more significant and moderate tohigh correlation coefficients intraregionally than interregion-ally
322 Qualitative Characters The frequency distribution forthe 16 qualitative characters of germplasm samples by regionsin Ethiopia and of the exotic accessions is shown in Supple-mental Table 5 For the purpose of avoiding redundancy andimproving readability the discrete characters are presented bygrouping related traits together
BudCushion andRelativeDegree of BudExtension Evaluationof the presence and absence of bud cushion revealed thatgreat proportion (70) of sugarcane landraces from Ethiopiado not have bud cushion Majority of the sample germplasmin all regions belong to this phenotypic class Approximatelyone-half (53) of the collection have their bud extendedabove the growth ring followed by those (41) having budtouching the growth ring
Relative Bud Shape Among the phenotypic classes for thischaracter tall deltoid was dominant (31) followed by ovate(17) and narrow ovate (12) While local accessions inmost regions had tall deltoid bud shape 50 of the intro-duced accessions belong to two phenotypic classes roundwith central germpore (30) and 20 having pentagonalbud shape Equal proportions (19) of the samples fromBenshangul-Gumz were Ovate and Ovate with emarginatebasal wing
Canopy Structure Relative Plant Erectness Colour of theLeaves and Colour of the Exposed Rind Out of the eightphenotypic classes observed compact tip droopy was themost frequent (35) Open semidroopy and open droopyclasses had fairly equal distribution With regard to relativeplant erectness seventy-five percent of the germplasm werefound to fall in twophenotypic classes erect (41) and almost
erect (34) for the whole region and country Most of theaccessions (90) showed light green or green leaf colourbut 20 from introduced collections were greenish yellowAmong phenotypic classes for colour of the exposed rindbrownish yellow yellow brown and yellowish green were themost frequent with fairly equal distribution
Relative Shape of Dewlap Double crescent had larger fre-quency (22) while squarish descending and flaring typeof dewlap shape had fairly equal proportions Seventeenpercent of germplasm from Tigray region had squarishdeltoid dewlap shape
Relative Degree of Internode Alignment and Internode ShapeForty-four percent of the sample germplasm were slightlyzigzag while there was almost equal frequency for straightand zigzag internode alignment for the entire region andcountry Straight or nearly straight (slightly zigzag) internodealignment of cane stalks is a very important character formechanized farming and postharvest handling in sugarcane[51] Forty percent of the collections were observed havingconcave-convex type of internode shape while 29 wascylindrical and 14 conoidal Other classes had fairly equalproportion
Type of Auricle and Relative Shape of Ligule An equalproportion of the germplasm totally constituting 84 ofthe collection belong to four phenotypic classes namelythose with no auricle and others with transitional and shortand lanceolate auricles Sixteen percent of the collectionsalso showed equal frequency for the remaining classesBenshangul-Gumz and Tigray regions had each equal 13accessions with falcate type of auricle Sixty-seven percentof the collections belong to three ligule shape classes cres-cent with lozenge (27) broad-crescent (21) and linear-crescent (19) The germplasm had almost equal proportionfor the other phenotypic classes of this character Nineteenpercent of the accessions fromBenshangul-Gumz region haddeltoid shape of ligule
Stalk Corky Cracks Corky Patches Growth Cracks and BudGroove Eighty-nine percent of the germplasm across regionshad no stalk corky cracks with only eleven percent exhibitingthis character All the germplasm in Gambella Harari andSomali had no stalk corky cracks On the other hand eighty-one percent of the germplasm for the whole region showedthe presence of stalk corky patches All the local germplasmin Benshangul-Gumz Gambella and Tigray had stalk corkypatches whereas 75 of the germplasm from Harari Region67 from Somali Region and 24 from Oromia Regionhad no stalk corky patches In the entire region 75 ofthe germplasm had no stalk growth cracks with only 25showing this character Sixty-three percent of the germplasmwere found to have bud groove Howevermost of the samplesin Harari (75) Amhara (63) and Oromia (53) Regionshad no bud groove
The results of this study indicated the wide distributionof phenotypic classes for the characters considered whichindicates the existence of different races and combination
Advances in Agriculture 15
of races in local genotypes of different regions in EthiopiaThis diversity of germplasmbased onphenotypicmarkers canhelp sugarcane breeders in identifying the identity of geno-types as well as maintaining genetic diversity Differencesin morphological characters of different sugarcane varietieshave been reported [52] Piscitelli [53] showed importantexomorphological qualitative characters of sugarcane varietythat are not influenced by environmental factors and thus canbe used as a selection tool in any breeding program
The amount of phenotypic diversity estimates based onthe ShannonndashWeaver diversity index (H1015840) are shown inTable 9 The 16 characters differed in their distribution aswell as the amount of variation Individual traits showed adifferent pattern of variation among accessions Estimatesof H1015840 for individual characters varied from 049 for stalkcorky cracks to 100 for bud groove with an overall meanof 080 Most of the characters were highly polymorphicwhereas relative plant erectness and presence or absenceof stalk corky patches scored moderate values The highdiversity values for the characters showed a wide variabilityamong genotypes According to [54] this index is also usedin genetic resource studies as a convenient measure of bothallelic richness and allelic evenness when using geneticdata
The highest diversity computed was on presence orabsence of bud groove with a diversity index of 100indicating that this character is very variable in the localsugarcane germplasm On the other hand the descriptorcomputed with the lowest diversity index value (049) wasthe presence or absence of stalk corky cracks indicatingthat this character is selected in the development of thegenotypes In agreement with the present study [55] workingon S officinarum accessions from the world collection ofsugarcane germplasm obtained a low index of 025 and 035for presence or absence of stalk corky patches and cracksrespectively
Their report also showed high diversity index values forinternode alignment growth cracks internode shape budcushion bud extension bud groove and ligule shape Sim-ilarly [56] experimenting on field collections of sugarcaneaccessions in Philippine reported high ShannonndashWeaverdiversity for stalk growth cracks bud cushion stalk corkypatch stalk corky cracks bud extension and auricle outershape Unlike the present study they have reported high indexfor stalk corky cracksMedium index values were reported forstalk internode shape and low index values for ligule shapeand bud shape High variability of sugarcane varieties fordewlap shape was also reported in [57]
The H1015840 pooled across characters by region ranged from079 in Oromiya to 085 in Tigray with an overall aver-age of 081 (Table 8) Most of the regions showed highShannonndashWeaver diversity index The populations that hadthe highest H1015840 were that of Tigray and introduction The H1015840averaged over regions for different characters was found torange from 044 for presence or absence of stalk corky cracksto 094 for relative degree of internode alignment and budgroove with an overall average of 081 (Table 9)
Assessment of the ShannonndashWeaver index also showedsensitivity to both the number of descriptor classes and the
distribution within phenotypic classes in each region Amongregions the mean values for the ShannonndashWeaver index arenot significantly different but differences were found whenindividual characters were considered These observationssuggest that some regions have high diversity for particulartraits while in other regions selective pressures might havereduced the variation to certain genotypes
4 Conclusions
The large number of landraces observed during the cur-rent study demonstrates the existence of diverse geneticresources of sugarcane in Ethiopia The landraces wouldserve to broaden the genetic base of the available sugarcanegermplasm Generally distribution of the overall phenotypicdiversity among sugarcane landraces in Ethiopia is unevenOur results showed that there was a wide range of variationthat existed in the local sugarcane landraces studied both atregional and within region levels Selection and differenti-ation of types might have taken place along a geographicalpattern perhaps associated with climate and use The presentstudy contributes to updating sugarcane descriptors adoptedfrom USDA-ARS as well as Bioversity passport data forthe future collection and evaluation As Ethiopian SugarCorporation is in the course of launching breeding programthe observed substantial variation of landraces would enablesugarcane breeders to design and practice breeding andselection programs to improve the crop
Data Availability
The data used to support the findings of this study areavailable from the corresponding author upon request
Conflicts of Interest
The authors declare that they have no conflicts of interest
Acknowledgments
The authors are grateful for the financial grant of SugarCorporation of Ethiopia Many thanks are due to EthiopianBiodiversity Institute (EBI) for its technical advice duringcollection of the local sugarcane genotypes Sincere thanksare forwarded to Ethiopian smallholder farmers who pre-served sugarcane germplasm and shared their knowledgeand experience Regional Zonal and District (ldquoWeredardquo)Agriculture Bureaus across Ethiopia are acknowledged fortheir support during the study
Supplementary Materials
These supplementary materials contain different tables sup-porting the results of the study These include passport dataof sugarcane germplasm collected and related informationon the materials local sugarcane germplasm collected andused formorphological diversity study introduced sugarcane
16 Advances in Agriculture
Table9Sh
anno
n-WeaverD
iversityIndices(H1015840)a
ndmeanandsta
ndarderror(plusmnS
E)of
land
racesineach
region
basedon
16discretecharacters
Characters
Amha
raBe
nsha
ngul-G
umz
Introd
uced
Oromia
SNNP
Tigray
ELb
Total
Presence
orabsenceo
fbud
cushion(BUDCUSH
ION)
083
095
095
089
082
098
090
088
Relativ
edegreeo
fbud
extension(BUDEX
TEND)
077
079
093
074
072
087
080
078
Relativ
ebud
Shape(BU
DSH
APE
)078
096
089
081
085
078
084
078
Canop
ystructure(C
ANOPY
)089
098
095
081
078
096
089
084
Relativ
eshape
ofdewlap(D
EWLA
PSHAP)
087
094
092
090
082
096
090
086
Relativ
eplant
erectness(ER
ECT)
080
082
050
067
071
088
073
064
Relativ
edegreeo
finterno
dealignm
ent(IN
ALIGN)
096
076
098
097
098
100
094
098
Relativ
einterno
deshape(IN
SHAPE
)086
069
095
077
083
091
083
082
Color
oftheleaves(LE
AFC
OLO
R)087
071
090
064
074
071
076
076
Type
ofou
tera
uricle(A
URI
CLE
OUT)
080
092
087
079
083
093
086
082
Relativ
eshape
ofligule(LIGSH
APE
)086
094
082
085
090
083
087
083
Presence
orabsenceo
fstalkcorkyc
racks(ST
ALK
CORK
C)
044
054
035
055
051
025
044
049
Presence
orabsenceo
fstalkcorkyp
atches(STA
LKCORK
P)066
000
072
079
059
087
060
069
Presence
orabsenceo
fstalkgrow
thcracks
(STA
LKCRA
CK)
066
100
088
062
088
099
084
080
Color
ofthee
xposed
rind(R
INDCOLE
)085
096
089
084
086
076
086
085
Presence
orabsenceo
fbud
groo
ve(BUDGRO
OVE)
095
100
072
100
100
099
094
100
Means
080plusmn0
03
081plusmn0
06
083plusmn0
04
079plusmn0
03
080plusmn0
03
085plusmn0
05
081plusmn0
03
080
b EL=
averageo
fentire
locatio
n
Advances in Agriculture 17
varieties used for morphological diversity study and fre-quency distribution for 16 qualitative characters in sugarcaneby regions in Ethiopia and country of sources of introducedgenotypes (Supplementary Materials)
References
[1] ldquoSpreading the word about clean solutions from sugarcanrdquoAccessed 04072018 httpwwwsugarcaneorg
[2] Central Statistics Agency (CSA) ldquoAgricultural sample surveyArea and production of major crops private peasant holdingsrdquoStatistical Bulletin Addis Ababa Ethiopia 2017
[3] T Esayas Exploration and Collection Characterization GeneticDiversity Analysis and Association of Traits for Yield and YieldComponents of Sugarcane (Saccharumspp) in Ethiopia [MasterThesis] Haramaya University Dire Dawa Ethiopia 2014
[4] J R Harlan Crops and Man American Society of AgronomyCrop Science Society of America Madison WI USA 2ndedition 1992
[5] R W Allard Principles of Plant Breeding JohnWilley and SonsNew York NY USA 2nd edition 1999
[6] B T Roach and J Daniels ldquoA review of the origin and improve-ment of sugarcanerdquo in Proceedings of Coper sugar InternationalSugarcane Workshop pp 1ndash32 1987
[7] P P Patil S S Patil and D U Patel ldquoGenetic diversity in sug-arcane (Saccharum spp Complex)rdquo Journal of Pharmacognosyand Phytochemistry vol 6 no 6 pp 570ndash573 2017
[8] R B Singh B Singh andR K Singh ldquoStudy of genetic diversityof sugarcane (Saccharum) species and commercial varietiesthrough TRAP molecular markersrdquo Indian Journal of PlantPhysiology vol 22 no 3 pp 332ndash338 2017
[9] R W Allard ldquoPopulation structure and sampling methodsrdquo inGenetic Resources in PlantsmdashTheir Exploration and Conserva-tion O H Frankel and E Bennett Eds IBP Handbook no 11pp 97ndash107 FA Davis Company Philadelphia PA USA 1970
[10] D R Marshall and A H D Brown ldquoOptimum samplingstrategies in genetic conservationrdquo inCrop Genetic Resources forToday and Tomorrow O H Frankel and J G Hawkes Eds pp53ndash80 Cambridge University Press 1975
[11] J G Hawkes Crop Genetic Resources Field Collection ManualPudoc Wagnningen Netherland 1980
[12] J G Hawkes The Diversity of Crop Plants Harvard UniversityPress London England 1983
[13] B P K Yerima Manual for Good Laboratory Practice Foodand Agriculture Organization of United Nation Addis AbabaEthiopia 1993
[14] A Walkley and I A Black ldquoAn examination of the degtjareffmethod for determining soil organic matter and a proposedmodification of the chromic acid titrationmethodrdquo Soil Sciencevol 37 no 1 pp 29ndash38 1934
[15] S R Olsen and L E Sommers ldquoPhosphorusrdquo in AgronomyMonograph A L Page et al Ed Methods of soil analysis part2 pp 403ndash430 ASA and SSSAMadisonWI USA 2nd edition1982
[16] R H Bray and L T Kurtz ldquoDetermination of total organic andavailable forms of phosphorus in soilsrdquo Soil Science vol 59 no1 pp 39ndash46 1945
[17] S Sahlemedhin andB Taye ldquoProcedures for soil and plant anal-ysisrdquo Tech Rep 74 National Soil Research Center EthiopianAgricultural Research Organization 2000
[18] F C Stickler S Weaden and AW Pauli ldquoLeaf area determina-tion in grain sorghumrdquo Journal of Agronomy and Crop Sciencevol 53 pp 187-188 1961
[19] Munsell Color ChartThe Munsell Book of Color Matte Collec-tion Munsell Color Chart NewWindsor NY USA 1990
[20] H O Hartley ldquoThe maximum F-ratio as a short cut test forheterogeneity of variancesrdquo Biometrika vol 37 pp 308ndash3121950
[21] L Pecetti P Annicchiarico and A B Damania ldquoBiodiversity ina germplasm collection of durumwheatrdquo Euphytica vol 60 no3 pp 229ndash238 1992
[22] L Pecetti and A B Damania ldquoGeographic variation intetraploid wheat (Triticum turgidum ssp turgidum convardurum) landraces from two provinces in Ethiopiardquo GeneticResources and Crop Evolution vol 43 no 5 pp 395ndash407 1996
[23] S K Jain C O Qualset G M Bhatt and K K Wu ldquoGeo-graphical patterns of phenotypic diversity in a world collectionof durumwheatsrdquoCrop Science vol 15 no 5 pp 700ndash704 1975
[24] K K Nkongolo L Chinthu M Malusi and Z Vokhiwa ldquoPar-ticipatory variety selection and characterization of sorghum(Sorghum bicolour (L) Moench) elite accessions from Malaw-ian gene pool using farmer and breeder knowledgerdquo AfricanJournal of Agricultural Research vol 3 pp 273ndash283 2008
[25] M B Semira ldquoInnovative methods and metrics for agricul-ture and nutrition actions soil quality and human nutritionin Ethiopiardquo Leverhulme Center for Integrative Research onAgriculture and Health (LCIRAH) 2017
[26] I A Khan A Khatri M A Siddiqui G S Nizamani and SRaza ldquoPerformance of promising sugarcane clone for yield andquality traits in different ecological zones of Sindhirdquo PakistanJournal of Botany vol 36 no 1 pp 83ndash92 2004
[27] M F Perera M E Arias D Costilla et al ldquoGenetic diversityassessment and genotype identification in sugarcane based onDNAmarkers andmorphological traitsrdquo Euphytica vol 185 no3 pp 491ndash510 2012
[28] C G Neil M Kay and C C Jack ldquoDiversity among mainlandUSA sugarcane cultivars examined by SSR genotypingrdquo Journalof the American Society of Sugar Cane Technologists vol 29 pp36ndash52 2009
[29] S SchenckMWCrepeauK KWu P HMooreQ Yu andRMing ldquoGenetic diversity and relationships in native HawaiianSaccharum officinarum sugarcanerdquo Journal of Heredity vol 95no 4 pp 327ndash331 2004
[30] R Gebretsadik H Shimelis M D Laing P Tongoona andN Mandefro ldquoA diagnostic appraisal of the sorghum farmingsystem and breeding priorities in Striga infested agro-ecologiesof Ethiopiardquo Agricultural Systems vol 123 pp 54ndash61 2014
[31] L E V Cesar A T Bruzi J A R Nunes et al ldquoGeneticdivergence of sugarcane varieties collected in the region ofLavras Minas Gerais Brazilrdquo Genetics and Molecular Researchvol 14 no 4 pp 14043ndash14049 2015
[32] C Grenier P J Bramel J A Dahlberg et al ldquoSorghums of theSudan analysis of regional diversity and distributionrdquo GeneticResources and Crop Evolution vol 51 no 5 pp 489ndash500 2004
[33] N V Nair R Balakrishnan and T V Sreenivasan ldquoVariabilityfor quantitative traits in exotic hybrid germplasm of sugarcanerdquoGenetic Resources and Crop Evolution vol 45 no 5 pp 459ndash464 1998
[34] R Nagarajan S Alarmelu and R M Shanthi ldquoStudies onvariation in interspecific hybrids of Saccharumrdquo Sugar Tech vol2 no 3 pp 42ndash46 2000
18 Advances in Agriculture
[35] D N Kamat and J R P Singh ldquoVariability in Sugarcane underrain fed conditionrdquo Sugar Technology Short Communicationsvol 3 no 1-2 pp 65ndash67 2001
[36] B S Rishipal A S M Chaudhary and S P Kadian ldquoStudieson character interrelationship in clonal generation of sugarcane(Saccharum complex hybrids)rdquo Indian Sugar vol 47 no 11 pp907ndash911 1998
[37] S Thippeswamy S T Kajjidoni P M Salimath and J V GoudldquoCorrelation and path analysis for cane yield juice quality andtheir component traits in sugarcanerdquo Short Communication vol5 no 1-2 pp 65ndash72 2003
[38] P S Verma S Pal and N K Karma ldquoGenetic variability andcorrelation studies in sugarcanerdquo Indian Sugar vol 49 pp 125ndash128 1999
[39] H A Esechie J W Maranville and W M Ross ldquoRelationshipof stalk morphology and chemical composition to lodgingresistance in sorghumrdquo Crop Science vol 17 no 4 p 609 1977
[40] P K Das B C Jena and N Nayak ldquoCharacter association andpath analysis of sugar yield in sugarcanerdquo Indian Sugar vol 46no 10 pp 805ndash808 1997
[41] K J Sreekumar M Kuriakose M Thomas D Alexander andS Santhakumari ldquoVariability heritability and correlation onstudies on the yield and quality characters of sugarcanerdquo IndianSugar vol 44 no 4 pp 243ndash249 1994
[42] J R Harlan J M de Wet and E G Price ldquoComparativeevolution of cerealsrdquo Evolution vol 27 no 2 pp 311ndash325 1973
[43] A Elings ldquoDurum wheat landraces from syria II patterns ofvariationrdquo Euphytica vol 54 no 3 pp 231ndash243 1991
[44] K Tato ldquoRainfall in Ethiopiardquo Ethiopian Geographical Journalvol 2 pp 28ndash36 1964
[45] A B L Stemler J R Harlan and J M J deWet ldquoThe sorghumsof ethiopiardquo Economic Botany vol 31 no 4 pp 446ndash460 1977
[46] P Annicchiarico L Pecetti and A B Damania ldquoRelationshipsbetween phenotypic variation and climatic factors at collectingsites in durum wheat landracesrdquo Hereditas vol 122 no 2 pp163ndash167 1995
[47] A B Damania L Pecetti C O Qualset and B O HumeidldquoDiversity and geographic distribution of adaptive traits inTriticum turgidum L (durum group) wheat landraces fromTurkeyrdquoGenetic Resources and Crop Evolution vol 43 no 5 pp409ndash422 1996
[48] A Ayana and E Bekele ldquoGeographical patterns of morpho-logical variation in sorghum (Sorghum bicolor (L) Moench)germplasm from Ethiopia and Eritrea quantitative charactersrdquoEuphytica vol 115 no 2 pp 91ndash104 2000
[49] D S Falconer Introduction to Quantitative Genetics JohnWilley and Sons Inc New York NY USA 3rd edition 1993
[50] R S Thorpe ldquoBiochemical analysis of geographical variationand racial affinitiesrdquo Biology Review vol 51 pp 407ndash452 1976
[51] M Farooq ldquoMorphological and agricultural characteristicsof sugarcane in relation to mechanized agriculture at NARCIslamabadrdquo Portu Rico Journal vol 1 pp 51ndash61 1989
[52] M Akhtar N N Elahi and M Ashraf ldquoEvaluation of exoticsugarcane varieties for agronomic characters and productivityrdquoPakistan Journal of Biological Sciences vol 4 no 1 pp 37ndash402001
[53] F R Piscitelli ldquoPrincipals caracteres exomorfologicos de cul-tures de cana de azucarrdquo TUC Iindustraly Agricos vol 7 pp 49ndash57 1994 (Spanish)
[54] A H D Brown and B S Weir ldquoMeasuring genetic variability inplant populationsrdquo in Isozyme in Plant Genetics Breeding S DTanksley and T J Orton Eds part A Elsevier Science 1983
[55] R Balakrishnan N V Nair and T V Sreenivasan ldquoA methodfor establishing a core collection of Saccharum officinarum Lgermplasm based on quantitative-morphological datardquo GeneticResources and Crop Evolution vol 47 no 1 pp 1ndash9 2000
[56] C A Nestor and B Q Reynaldo Sugarcane Variety Improve-ment in Southeast Asia and the Pacific for Enhanced andSustainable Productivity ndash Germplasm and Disease IndexingComponents Institute of Plant Breeding (IPB) College ofAgriculture (CA) UP Los Banos (UPLB) 2003
[57] A Maqbool J Muhammad and A Sagheer ldquoAgronomic traitsand morphological characteristics of some exotic varieties ofsugarcanerdquoPakistan Journal of Agricultural Research vol 19 no4 2006
Figure 1 A map of Ethiopia showing sugarcane germplasm collection sites (black circles)
Table 1 Location information of the test sites
Location Latitude Longitude Altitude (masl)119886 Rainfall (mm) Temperature (∘C)Minimum Maximum
Wonji 8∘311015840 N 39∘121015840 E 1550 800 153∘C 269∘CMetehara 8∘511015840 N 39∘ 521015840 E 950 554 175∘C 326∘C119886masl = meters above sea levelSource meteorological Stations of respective locations
22 Diversity Study
221 Plant Materials A total of 211 sugarcane (Saccharumspp) accessions consisting of 196 landraces (SupplementalTable 3) and 15 introduced commercial varieties including2 standard varieties (Supplemental Table 4) were used forthis study The landraces represent collections from allgeographical regions across Ethiopia maintained at fieldconservation garden at Wonji and Metehara Sugar EstatesSugar Corporation of Ethiopia Sampling of the landraces wasmade on representation basis stratified systematic samplingmethod to a given range of geographic area altitudinalranges and a range of morphological traits The introducedmaterials were commercial varieties under production indifferent estates where the two were standard varieties
The 196 landraces were collections from the followingregions of Ethiopia Amhara (47) Benshangul-Gumz (10)Gambella (3) Harari (2) Oromiya (65) SNNPR (59) Somali(3) and Tigray (7) (Figure 1) The altitude of the collectionsites for the landraces used in this study ranged from 454to 2687 meters above sea level representing the distributionof the crop in Ethiopia The introduced materials were fromBarbados (6) Cuba (1) India (5) Mexico (1) and SouthAfrica (2)
222Methods The plant materials were grown at Wonji andMetehara Sugar Estates of Sugar Corporation of Ethiopiaduring the 201213 growing season Details of the plantinglocations are shown in Table 1 Each accession was grown ina single row plot of 5m long and 145m between rows and20 cm between plants within a row with two replications inrandomized complete block design
Uniform crop management practices were applied asrecommended for the areas Urea was applied 25 monthsafter planting at a rate of 200 kgsdothaminus1 atWonji and 400 kgsdothaminus1at Metehara
Accessions from regions with sample size less than 12were included in adjacent regions to reduce experimentalerror due to small sample size Hence the two Harari andthree Somali accessions were included in Tigray and thethree Gambella accessions were included in Benshangul-GumzThis reduced the 8 regions of Ethiopia fromwhich thelandraces were originally drawn to five With the introducedmaterials included 6 regions of origin were used in thestatistical analyses
Data on 17 quantitative stalk characters was recordednamely sprout count 1 and 2 months after planting(SPC1MAP and SPC2MAP) tiller counts 4 and 5month afterplanting (TC4MAP and TC5MAP) stalk count 10 months
4 Advances in Agriculture
after planting (STC10MAP) hand refractometer brix reading10 months after planting (HRBrix10MAP) millable stalkcount per hectare (MSCHA) single cane weight (SCW)number of internode (NOI) internode length (IL) stalkheight (SH) stalk diameter (SD) leaf length (LL) leaf width(LW) leaf area (LA) cane yield quintal per hectare (CYHA)and sugar yield quintal per hectare (SY) Data on 4 juicequality parameters ie brix percent (Brix) pol percent(pol ) purity percent (purity ) and sugar percent (SR) was also recorded For every accession ten plants wereused for recording data for quantitative characters whichwere recorded on plot basis Count data and cane yield wererecorded considering all cane stalks from the whole plot Forquantitative leaf characteristics measurement a proceduredeveloped in [18] was used
To categorize each accession morphologically sugarcanedescriptors adopted fromUSDA-ARSwere employed (GRIN2004) Data on 16 qualitative traits was recorded namelypresence or absence of bud cushion (BUDCUSHION) rel-ative degree of bud extension (BUDEXTEND) relative budshape (BUDSHAPE) relative shape of dewlap (DEWLAP-SHAP) type of outer auricle (AURICLEOUT) presence orabsence of stalk corky cracks (STALKCORKC) presenceor absence of stalk corky patches (STALKCORKP) relativeshape of ligule (LIGSHAPE) presence or absence of stalkgrowth cracks (STALKCRACK) presence or absence of budgroove (BUDGROOVE) relative plant erectness (ERECT)relative degree of internode alignment (INALIGN) relativeinternode shape (INSHAPE) colour of the leaves (LEAF-COLOR) colour of the exposed rind (RINDCOLE) andcanopy structure (CANOPY) Each accession was scored forthe most frequent character-state Leaf colour and colourof the exposed rind were examined and scored using theMunsell colour chart [19]
223 Statistical Analyses First analysis of variance wasmadefor the 21 characters for each location Homogeneity ofthe error variances among the locations was assessed in[20] F-max method for each of the 21 characters The testestablished the homogeneity of the error variances for allcharacters except TC4MAP For TC4MAP logarithmic datatransformation which is recommendable for continuousdata was used to homogenize the error variance Thendata of all the characters were subjected to test location-accession-related analysis of variance to determine effects oftest sites and genotype X environment (G X E) interactionThe results showed the nonsignificance of effects of testsites for SPC1MAP IL LW Pol sugar percent (SR) andthe nonsignificance of G X E interactions for SPC2MAPSCW Brix Pol and Brix10MAP For the other charactersboth test site effects and G X E interactions were significant(Plt005)
Analysis of variance was made for 21 quantitative char-acters following the procedure used in [21 22] The meansquares of the regions were tested against pooled meansquares of accessions within regions The pooled meanssquares for accessions within regions of origin and the meansquares of accessions within each region were tested against
the pooled within-region error mean squares Means rangesfor means and percent coefficients of variation for all thecharacters were computed for each region of origin and forthe entire data The regional means were compared usingDuncanrsquos multiple range testing Correlations between thecharacters were computed at three levels First correlationsof the characters were assessed based on the 211 accessionmeans Then interregion correlation was computed usingthe means of characters for each region Finally a series ofintraregion correlation coefficient matrices were obtained foreach region using the accession means from that region forthe characters
For qualitative characters phenotypic frequency distri-butions were worked out for all the sample germplasm andlocations The ShannonndashWeaver diversity index (H1015840) wascomputed using the phenotypic frequencies to assess thephenotypic diversity for each character for all accessions TheShannonndashWeaver diversity index as described in [23] is givenas
1198671015840 = minus119899
sum119894=1
119901119894log119890(119901119894) (1)
where pi is the proportion of accessions in the 119894th class of ann-class character and n is the number of phenotypic classesof traits Each 1198671015840 value was divided by its maximum value(logen) and normalized in order to keep the values between0 and 1 By pooling various characters across the regions theadditive properties of 1198671015840 were used to evaluate diversity ofregions and characters within the population The averagediversity index (H1015840) over n traits was estimated as H1015840 =sumH1015840n
3 Results and Discussion
31 Collection of Sugarcane Germplasm In Ethiopia thehistory of sugarcane cultivation by smallholder farmers pre-ceded that of commercial cultivation As documented in thehistory of a monastery in Northern Ethiopia sugarcane hasbeen grown in the country since the 16th century [2] In thisstudy local sugarcane germplasm exploration and collectionwere conducted all across Ethiopia in the regional statesMore than 300 local sugarcane genotypes were collectedduring 20102011 and passport data of the genotypes ispresented in Supplemental Table 1 The collected germplasmwere planted at five locations across the country with respectto their collection area These are Wondogenet AgriculturalResearchCenter JimmaAgricultural ResearchCenterMechaWereda Agricultural Bureau nursery field at Picolo AbaySirinka Agricultural Research Center (at Kobo Subcenter)and research field of Haramaya University at Diredawa(Tonny Farm) The clones have been monitored during the201112 season for symptoms of major diseases and insectsThese clones were transferred to commercial sugarcane plan-tation estates at Wonji and Metehara for further selectionand maintenance No major diseases and insect pests weredetected in all the germplasm collected except sometimestermites and borer were observed (Supplemental Table 2)
Advances in Agriculture 5
Table 2 Mean soil physical and chemical properties of sugarcane fields in the surveyed regions
Regions Soil chemical properties Texture ()PH EC(dsm) OC N P (ppm) K(ppm) Sand Silt Clay
Most of the sampled germplasm had acceptable levels of juicerefractometer readings (expressed in degree brix) relative tothe history of the ages of the samples (Supplemental Table 1)The number of harvestablesaleable cane stalks obtained inplant cane and ratoon crops and the number of harvestableratoon crops showed variation region to region and zone tozone and sometimes district (ldquoweredardquo)-wise There is alsovariation in maturity time of the clones both in plant caneand in ratoon crops (Supplemental Table 2)These are mainlydepending on the variety soil condition cultural practicesand climatic conditions
The genotypes collected from the respective zones inthe study regions were recorded by their local namesSometimes similar varieties bear different names at differentlocalities and vice versa Farmers mentioned a broad rangeof sugarcane landraces that had been grown in the areasand maintained for generations Some of the landraces werecommonly recognized by most farmers within and acrosszones of the respective regions whereas some were rarevarieties known only by few farmers The large number oflandraces observed during the current study demonstratesthe existence of diverse genetic resources of sugarcane inEthiopia The diversity could have evolved presumably due todiverse climatic conditions low input management systemshigh pests and disease pressure and continuous selectionby farmers Smallholder farmers face diverse environmentstresses and have multiple production objectives that affectselection of genotypes [24] There were no formally releasedimproved sugarcane varieties grown by farmers in the studyregions except in few places This was mainly observedin SNNPR at Wolayta and Kembata-Tembaro zones Thesevarieties have been informally introduced to these areas byseasonal workers employed at the sugar estates especially atWonji and Metehara
The collected landraces would serve to broaden thegenetic base of the available sugarcane germplasm Thesugar industry in Ethiopia is currently establishing sugarcanebreeding programTherefore collection and efficient charac-terization of germplasm are vital to strengthen the breedingprogram
311 Physical and Chemical Properties of Soils of Sugar-cane Germplasm Collection Areas Table 2 summarizes thephysical and chemical properties of soils in the sugarcane
production fields of surveyed areas Sugarcane productionby smallholder farmers in the surveyed areas is conductedacross the seasons where harvesting and planting are done aslong as there is available moisture Therefore the soil samplesrepresent the sugarcane production season and the altitudinalranges of the respective areas
Generally the organic carbon content of soils in sugarcaneproducing areas across the country is low to very low (2-4lt2) Amhara Gambella SNNP Somali and Tigray Regionsare in a very low category (lt2) Similarly across regions thetotal N content is low except Harari and Oromia regionswhere the average total N is medium This problem shouldbe addressed either with augmenting the soil with inorganicfertilizers or through appropriate management practices likecrop residue management Available phosphorus is high inAmhara Gambella and SNNP Regions whereas moderatevalues were observed in Harari Oromia and Tigray RegionsGenerally the level of phosphorus in soils of sugarcaneproducing areas in these regions is adequate In Benshangul-Gumz and Somali Regions low level of phosphorus wasrecorded which should be addressed with due attentionAvailable K is sufficient for normal sugarcane growth in soilsof all regions Relatively lower values of K were recorded insoils of Harari and Tigray Regions It should be noted thatorganic carbon EC PH and variability of P andK are affectedby rainfall and temperature that are seasonal and the valuesrecorded refer to the seasonwhen the soil samples were takenGenerally Ethiopian soils are deficient in various essentialnutrients like boron nitrogen phosphorus potassium sulfurzinc and copper although severity differs from region toregion [25] Particularly the loss of P and N resulting fromthe use of dung and crop residues for fuel is demonstrated tobe equivalent to the total amount of commercial fertilizer use
Across regions in Ethiopia generally clay type of soildominates followed by silt except in Gambella Region wherethe dominant soil texture was sand Soil management optionsrelated to clay soil may address soil related problems insmallholder sugarcane growing areas across Ethiopia
32 Diversity of Sugarcane Landraces
321 Quantitative Characters
Univariate Statistics Analysis of variance depicted highlysignificant differences (p lt 001) between accessions pooled
6 Advances in Agriculture
over the regions for the 21 characters of the 211 sugarcaneaccessions studied and significant difference for the 6 regionsof origin for 6 characters (Table 3) The results suggestedthe occurrence of significant phenotypic variation betweenthe accessions as a whole Significant variations of differentsugarcane stalk and juice quality characters was also reportedin similar studies conducted elsewhere [26 27] Region-wise partitioning of the variance indicated significant within-region differences (p lt 005) among the populations withinalmost all regions for the characters NOI IL SH SD LLLW LA Brix Pol Purity SR MSCHA SCW CYHAand SY for 8 characters within Tigray for TC4MAP andTC5MAP within Oromiya and SNNPR Brix10MAP showedno significant difference within all regions
In general within-region variation was greater for stalkdiameter single cane weight millable stalk count and caneand sugar yield than for other characters for all the regionsAssuming that a significant portion of the phenotypic vari-ation is genetic it would be possible to make selection forany of this group of characters within a particular regionIt was apparent that the variance between accessions pooledover regions was greater than between regions Therefore inorder to sample the variation effectively it would be necessaryto sample the variable populations from different localitiesin a region The study in [28] on genetic diversity amongmain land USA sugarcane cultivars showed high geneticdiversitywithin populations of different regions than betweenpopulations
Duncanrsquos multiple range testing for regional means for allthe characters is shown in Table 4 Relatively much differenti-ationwas observed for stemdiameter and juice purity percentcompared to other parameters Important yield componentslike MSCHA SCW SH and SD have shown higher valuesin Tigray SNNPR and Oromiya regions High cane andsugar yield were recorded for accessions in Tigray regionfollowed by SNNPR Oromiya and introduction which werestatistically at par Statistically similar lower values of caneand sugar yield were recorded in Amhara and Benshangulcollections
Higher value of sucrose percent in cane was also observedfor accessions from Tigray and the lowest value was inAmhara accessions while other regions were statisticallysimilar Generally for most of the characters high valuewas obtained from accessions in Tigray region followed bySNNPROromiya and introduction Therefore the materialsfrom these regions can be used for selection of cane and sugaryield per se and in breeding program The local accessions inmany of the regions showed superiority over the introducedcommercial varieties currently in production for most of thecharacters including cane and sugar yield (Table 4)This maybe due to the adaptation and coevolution of the materialswith different stresses for long period of time in the countryrsquosagroecology These locally adapted genes conferring yieldadvantage could be harnessed through strategic crossing andselection for improvement of the crop Superiority of locallandraces over introduced varieties was reported [29 30]
The range of variation of the accession means establishedwide variation between the regions and the accessions withinthe regions for the characters studied (Table 5) Accordingly
the maximum score was 41 times the minimum for sproutcount one month after planting 32 for sprout count twomonths after planting 25 for tiller count four months afterplanting 22 for sugar yield 20 for cane yield 12 for millablestalk count 8 for stalk count 10 months after planting 7for single cane weight 6 for tiller count five months afterplanting and 2 for other yield components and sugar qual-ity parameters except hand refractometer brix reading tenmonths after planting and juice purity percent which scored1 The same trend was observed between accessions withina particular region However accessions within AmharaBenshangul-Gumz and introduction showed wider ranges ofvariation than those accessions within other regions for themajority of the characters The wide range of geographic andclimatic features of the regions in Ethiopia and the originaldifferentiation and further farmers selection for productionniches and uses for hundreds of years may have resultedin the accessions to possess a tremendously high degree ofmorphological variation The reason for introduced varietiesshowing high variation also could be due to broader geo-graphic spectrum fromwhere theywere initially acquired anddifference in their parental source and genealogical historyIt was demonstrated even with these limited numbers ofsamples (15 genotypes) that introduction of genetic materialshas practically broadened the genetic base of the sugarcanegermplasm for the breeding program
Coefficient of variation measured as the ratio of standarddeviation to the corresponding overall mean and expressedas percentage is useful to compare different charactersmeasured in different units It is also useful to compare samecharacter in different groups of populations with differentsample size mean and variance or different characters indifferent populations In the current study high coefficientsof variation were observed between regions and within eachregion for sprout as well as tiller counts in the monthsthe data was recorded stalk count 10 months after plant-ing and millable stalk count at harvest cane and sugaryield single cane weight and leaf area (Table 6) Thoughaccessions from a particular region were more variable fora specific character compared to other regions accessionsfromAmhara Benshangul-Gumz introduction and SNNPRwere more variable than accessions from other regions Thisresult demonstrated the tremendous variability of sugar-cane germplasm from these regions The accessions fromTigray and Oromiya had low coefficients of variation formany characters indicating relatively high within-regionuniformity The high coefficients of variation observed formost of the characters agreed well with those reportedin [31] for indigenous sugarcane of Brazil Similar resultswere also reported in sorghum [32] The different levels ofregional variability of a particular character could be due todifferences in forces of selection andor differences in theintensity of a particular selecting force It could also be dueto random effect and reduced sample parents of parentalmaterials moved differently by various human populationmigration and the interaction of these with Darwinianforces
Results of this study suggested the presence of wide rangeof variations of characters determined by univariate statistics
This is in agreement with previous studies on sugarcanegermplasm [33ndash35]
This is the first study to report information on geo-graphical pattern of variation in Ethiopian local sugarcanegermplasm which has been lacking so far Our results pro-vided experimental evidence on occurrence of geographicalvariation and significant within-region variation where it washigh in the regions such as Amhara Benshangul-Gumz andSNNPR
The overall patterns of similarity or difference betweenregions seemed to depend on environmental factors such asrainfall temperature length of growing season and altitudeWide variability of millable stalk count at harvest singlecane weight and plant height was observed among regionswhere these characters demonstrated high correlation withcane and sugar yield (Table 7) Therefore breeders can utilizeaccessions of regions showing variability for these charactersin selection programs and to design breeding strategies toproduce varieties with best commercial merits
Bivariate Statistics Correlation coefficients worked out onthe 21 quantitative characters are shown for the entire data(Table 7) between regions (Table 8) and within regions(data not shown) Results of the association studies for theentire data showed that cane and sugar yield had highlysignificant positive correlation with most of the quantitativestalk characters and juice quality parameters However theassociation of tiller counts 4 and 5 months after plantingstalk count 10 months after planting millable stalk countat harvest single cane weight and plant height with caneand sugar yield was stronger suggesting these componentcharacters asmain contributing factor to cane and sugar yieldIn fact sugar yield is the product of cane yield and sucrose per-cent cane surely being influenced with components affectingcane yield Similar trend was reported by earlier sugarcaneworkers [36 37] In sugarcane the cane and sugar yields areconsidered to be the complex characters The information onthe phenotypic interrelationship of cane yield and sugar yieldwith their component character per se would be of immensehelp to sugarcane breeders
Stem diameter had significant positive association withboth cane and sugar yield indicating the significance ofthis trait in improving both cane and sugar yields Similarreports have beenmade by earlier sugarcane workers [37 38]Stalk diameter was also positively and significantly correlatedwith plant height It is one of the traits that are related tolodging resistance [39] and the positive associations withheight would help in reducing the chance of lodging as heightincreases Among the physiological attributes included in thestudy leaf area had highly significant positive associationwith cane yield Similar observation was also made in [37]Sucrose percent which is an important juice quality attributehad highly significant positive association with sugar yieldand simultaneously cane yield also had highly significantassociation with sugar yield Based on the magnitude ofcorrelation coefficient values it can be said that cane yield ismuch more important than sucrose percent in determiningthe sugar yield and the same has also been reported by othersugarcane workers [37 40] Similar to observations made in
[37 41] results of the current study showed that all the qualityparameters like sucrose percent pol percent brix percentand purity percent were significantly correlated in positivedirection This indicated any of these juice quality traitscould be considered for selection leading to the simultaneousimprovement in the remaining quality traits
Ten of the 21 characters also showed significant positivecorrelations with altitude of the collection sites (Table 7) Asdescribed in [42] ecological characteristics have influencedthe genotypic constitution of landraces during domesticationand hence a relationship exists between the agroecology inthe collection site and the morphological characteristics ofthe landraces Thus positive correlation between collectionsite variables and plant characteristics would imply that thevariation between accessionsmay be related to agroecologicalvariations among the collection sites [43]
The correlation coefficient between number of internodesand altitude was negative and significant indicating thatother environmental factors (other than altitude) andornonenvironmental factorsmight account for the variation forthis particular character Since temperature decreases withan increase of altitude in Ethiopia [44 45] it is more likelythat temperature has exerted strong selection pressure innumber of internodes in the genotypes Information on therelationships between environmental factors of the collect-ing sites and morphophysiological variation of germplasmcould enhance the understanding of evolutionary adaptivepatterns which could assist germplasm collectors and users[46]
Character associations between region and within regionalso followed similar fashion like that of the entire dataNumber of sprout tiller and stalk counts and sugar qualityparameters showed significant positive correlations with caneand sugar yield
Though not significant and strong the correlation of leaflength and width and leaf area (leaf area = leaf length Xleaf width X 075) with cane and sugar yield was negativein the interregion correlations This kind of relationship wasalso exhibited between these leaf characteristics and yieldcomponents like tiller and stalks counts plant height andsingle cane weight and stalk diameter and sugar qualityparameters This may have been attributed to the variationof temperature with altitude and other environmental factorsacross the regions resulting in such type of relationshipSuch relationship also calls for the need to have enough dataon environmental variables of the collection sites such astemperature and rainfall if germplasm collection is requiredtomeet its objectives effectively and efficiently Unfortunatelydata on such useful environmental variables like temperatureand rainfall are lacking in the passport data of the currentEthiopian sugarcane germplasm collections The significanceof environmental factors of the collection sites as veryimportant determinants in structuring morphophysiolog-ical variations was reported in tetraploid wheat [22 4748]
Association of characters among yield its componentsand other economic traits is important for the interpretationof the patterns of variation and making selection in breed-ing program and combining several desirable attributes It
suggests the advantage of a scheme of selection for morethan one character at a time The correlation between char-acters may arise from linkage or from developmental geneticinteractions with orwithout a purely phenotypic componentIt could arise also due to differential phenotypic plasticityof characters themselves Within the limit of experimentalerror and environmental effects high correlation coefficientsbetween characters may show that the characters sharesome common element of genetic control (ie pleiotropylinkage) between genes or else from independently controlledcharacters responding similarly to geographic variation inselection pressures [49] The interregion correlation coef-ficient between the characters measures the consistency oftheir patterns of regional variation while the intraregioncorrelation coefficient measures the association arising fromgenetic factors but is not affected by regional variation[50] Since this study showed significant positive correlationsintraregionally for some character combinations it wouldseem that common genetic control might be playing a role inbringing about correlations between the various charactersIt appeared that different response to regional variationwas playing a greater role than different genetic controlas shown by the many more significant and moderate tohigh correlation coefficients intraregionally than interregion-ally
322 Qualitative Characters The frequency distribution forthe 16 qualitative characters of germplasm samples by regionsin Ethiopia and of the exotic accessions is shown in Supple-mental Table 5 For the purpose of avoiding redundancy andimproving readability the discrete characters are presented bygrouping related traits together
BudCushion andRelativeDegree of BudExtension Evaluationof the presence and absence of bud cushion revealed thatgreat proportion (70) of sugarcane landraces from Ethiopiado not have bud cushion Majority of the sample germplasmin all regions belong to this phenotypic class Approximatelyone-half (53) of the collection have their bud extendedabove the growth ring followed by those (41) having budtouching the growth ring
Relative Bud Shape Among the phenotypic classes for thischaracter tall deltoid was dominant (31) followed by ovate(17) and narrow ovate (12) While local accessions inmost regions had tall deltoid bud shape 50 of the intro-duced accessions belong to two phenotypic classes roundwith central germpore (30) and 20 having pentagonalbud shape Equal proportions (19) of the samples fromBenshangul-Gumz were Ovate and Ovate with emarginatebasal wing
Canopy Structure Relative Plant Erectness Colour of theLeaves and Colour of the Exposed Rind Out of the eightphenotypic classes observed compact tip droopy was themost frequent (35) Open semidroopy and open droopyclasses had fairly equal distribution With regard to relativeplant erectness seventy-five percent of the germplasm werefound to fall in twophenotypic classes erect (41) and almost
erect (34) for the whole region and country Most of theaccessions (90) showed light green or green leaf colourbut 20 from introduced collections were greenish yellowAmong phenotypic classes for colour of the exposed rindbrownish yellow yellow brown and yellowish green were themost frequent with fairly equal distribution
Relative Shape of Dewlap Double crescent had larger fre-quency (22) while squarish descending and flaring typeof dewlap shape had fairly equal proportions Seventeenpercent of germplasm from Tigray region had squarishdeltoid dewlap shape
Relative Degree of Internode Alignment and Internode ShapeForty-four percent of the sample germplasm were slightlyzigzag while there was almost equal frequency for straightand zigzag internode alignment for the entire region andcountry Straight or nearly straight (slightly zigzag) internodealignment of cane stalks is a very important character formechanized farming and postharvest handling in sugarcane[51] Forty percent of the collections were observed havingconcave-convex type of internode shape while 29 wascylindrical and 14 conoidal Other classes had fairly equalproportion
Type of Auricle and Relative Shape of Ligule An equalproportion of the germplasm totally constituting 84 ofthe collection belong to four phenotypic classes namelythose with no auricle and others with transitional and shortand lanceolate auricles Sixteen percent of the collectionsalso showed equal frequency for the remaining classesBenshangul-Gumz and Tigray regions had each equal 13accessions with falcate type of auricle Sixty-seven percentof the collections belong to three ligule shape classes cres-cent with lozenge (27) broad-crescent (21) and linear-crescent (19) The germplasm had almost equal proportionfor the other phenotypic classes of this character Nineteenpercent of the accessions fromBenshangul-Gumz region haddeltoid shape of ligule
Stalk Corky Cracks Corky Patches Growth Cracks and BudGroove Eighty-nine percent of the germplasm across regionshad no stalk corky cracks with only eleven percent exhibitingthis character All the germplasm in Gambella Harari andSomali had no stalk corky cracks On the other hand eighty-one percent of the germplasm for the whole region showedthe presence of stalk corky patches All the local germplasmin Benshangul-Gumz Gambella and Tigray had stalk corkypatches whereas 75 of the germplasm from Harari Region67 from Somali Region and 24 from Oromia Regionhad no stalk corky patches In the entire region 75 ofthe germplasm had no stalk growth cracks with only 25showing this character Sixty-three percent of the germplasmwere found to have bud groove Howevermost of the samplesin Harari (75) Amhara (63) and Oromia (53) Regionshad no bud groove
The results of this study indicated the wide distributionof phenotypic classes for the characters considered whichindicates the existence of different races and combination
Advances in Agriculture 15
of races in local genotypes of different regions in EthiopiaThis diversity of germplasmbased onphenotypicmarkers canhelp sugarcane breeders in identifying the identity of geno-types as well as maintaining genetic diversity Differencesin morphological characters of different sugarcane varietieshave been reported [52] Piscitelli [53] showed importantexomorphological qualitative characters of sugarcane varietythat are not influenced by environmental factors and thus canbe used as a selection tool in any breeding program
The amount of phenotypic diversity estimates based onthe ShannonndashWeaver diversity index (H1015840) are shown inTable 9 The 16 characters differed in their distribution aswell as the amount of variation Individual traits showed adifferent pattern of variation among accessions Estimatesof H1015840 for individual characters varied from 049 for stalkcorky cracks to 100 for bud groove with an overall meanof 080 Most of the characters were highly polymorphicwhereas relative plant erectness and presence or absenceof stalk corky patches scored moderate values The highdiversity values for the characters showed a wide variabilityamong genotypes According to [54] this index is also usedin genetic resource studies as a convenient measure of bothallelic richness and allelic evenness when using geneticdata
The highest diversity computed was on presence orabsence of bud groove with a diversity index of 100indicating that this character is very variable in the localsugarcane germplasm On the other hand the descriptorcomputed with the lowest diversity index value (049) wasthe presence or absence of stalk corky cracks indicatingthat this character is selected in the development of thegenotypes In agreement with the present study [55] workingon S officinarum accessions from the world collection ofsugarcane germplasm obtained a low index of 025 and 035for presence or absence of stalk corky patches and cracksrespectively
Their report also showed high diversity index values forinternode alignment growth cracks internode shape budcushion bud extension bud groove and ligule shape Sim-ilarly [56] experimenting on field collections of sugarcaneaccessions in Philippine reported high ShannonndashWeaverdiversity for stalk growth cracks bud cushion stalk corkypatch stalk corky cracks bud extension and auricle outershape Unlike the present study they have reported high indexfor stalk corky cracksMedium index values were reported forstalk internode shape and low index values for ligule shapeand bud shape High variability of sugarcane varieties fordewlap shape was also reported in [57]
The H1015840 pooled across characters by region ranged from079 in Oromiya to 085 in Tigray with an overall aver-age of 081 (Table 8) Most of the regions showed highShannonndashWeaver diversity index The populations that hadthe highest H1015840 were that of Tigray and introduction The H1015840averaged over regions for different characters was found torange from 044 for presence or absence of stalk corky cracksto 094 for relative degree of internode alignment and budgroove with an overall average of 081 (Table 9)
Assessment of the ShannonndashWeaver index also showedsensitivity to both the number of descriptor classes and the
distribution within phenotypic classes in each region Amongregions the mean values for the ShannonndashWeaver index arenot significantly different but differences were found whenindividual characters were considered These observationssuggest that some regions have high diversity for particulartraits while in other regions selective pressures might havereduced the variation to certain genotypes
4 Conclusions
The large number of landraces observed during the cur-rent study demonstrates the existence of diverse geneticresources of sugarcane in Ethiopia The landraces wouldserve to broaden the genetic base of the available sugarcanegermplasm Generally distribution of the overall phenotypicdiversity among sugarcane landraces in Ethiopia is unevenOur results showed that there was a wide range of variationthat existed in the local sugarcane landraces studied both atregional and within region levels Selection and differenti-ation of types might have taken place along a geographicalpattern perhaps associated with climate and use The presentstudy contributes to updating sugarcane descriptors adoptedfrom USDA-ARS as well as Bioversity passport data forthe future collection and evaluation As Ethiopian SugarCorporation is in the course of launching breeding programthe observed substantial variation of landraces would enablesugarcane breeders to design and practice breeding andselection programs to improve the crop
Data Availability
The data used to support the findings of this study areavailable from the corresponding author upon request
Conflicts of Interest
The authors declare that they have no conflicts of interest
Acknowledgments
The authors are grateful for the financial grant of SugarCorporation of Ethiopia Many thanks are due to EthiopianBiodiversity Institute (EBI) for its technical advice duringcollection of the local sugarcane genotypes Sincere thanksare forwarded to Ethiopian smallholder farmers who pre-served sugarcane germplasm and shared their knowledgeand experience Regional Zonal and District (ldquoWeredardquo)Agriculture Bureaus across Ethiopia are acknowledged fortheir support during the study
Supplementary Materials
These supplementary materials contain different tables sup-porting the results of the study These include passport dataof sugarcane germplasm collected and related informationon the materials local sugarcane germplasm collected andused formorphological diversity study introduced sugarcane
16 Advances in Agriculture
Table9Sh
anno
n-WeaverD
iversityIndices(H1015840)a
ndmeanandsta
ndarderror(plusmnS
E)of
land
racesineach
region
basedon
16discretecharacters
Characters
Amha
raBe
nsha
ngul-G
umz
Introd
uced
Oromia
SNNP
Tigray
ELb
Total
Presence
orabsenceo
fbud
cushion(BUDCUSH
ION)
083
095
095
089
082
098
090
088
Relativ
edegreeo
fbud
extension(BUDEX
TEND)
077
079
093
074
072
087
080
078
Relativ
ebud
Shape(BU
DSH
APE
)078
096
089
081
085
078
084
078
Canop
ystructure(C
ANOPY
)089
098
095
081
078
096
089
084
Relativ
eshape
ofdewlap(D
EWLA
PSHAP)
087
094
092
090
082
096
090
086
Relativ
eplant
erectness(ER
ECT)
080
082
050
067
071
088
073
064
Relativ
edegreeo
finterno
dealignm
ent(IN
ALIGN)
096
076
098
097
098
100
094
098
Relativ
einterno
deshape(IN
SHAPE
)086
069
095
077
083
091
083
082
Color
oftheleaves(LE
AFC
OLO
R)087
071
090
064
074
071
076
076
Type
ofou
tera
uricle(A
URI
CLE
OUT)
080
092
087
079
083
093
086
082
Relativ
eshape
ofligule(LIGSH
APE
)086
094
082
085
090
083
087
083
Presence
orabsenceo
fstalkcorkyc
racks(ST
ALK
CORK
C)
044
054
035
055
051
025
044
049
Presence
orabsenceo
fstalkcorkyp
atches(STA
LKCORK
P)066
000
072
079
059
087
060
069
Presence
orabsenceo
fstalkgrow
thcracks
(STA
LKCRA
CK)
066
100
088
062
088
099
084
080
Color
ofthee
xposed
rind(R
INDCOLE
)085
096
089
084
086
076
086
085
Presence
orabsenceo
fbud
groo
ve(BUDGRO
OVE)
095
100
072
100
100
099
094
100
Means
080plusmn0
03
081plusmn0
06
083plusmn0
04
079plusmn0
03
080plusmn0
03
085plusmn0
05
081plusmn0
03
080
b EL=
averageo
fentire
locatio
n
Advances in Agriculture 17
varieties used for morphological diversity study and fre-quency distribution for 16 qualitative characters in sugarcaneby regions in Ethiopia and country of sources of introducedgenotypes (Supplementary Materials)
References
[1] ldquoSpreading the word about clean solutions from sugarcanrdquoAccessed 04072018 httpwwwsugarcaneorg
[2] Central Statistics Agency (CSA) ldquoAgricultural sample surveyArea and production of major crops private peasant holdingsrdquoStatistical Bulletin Addis Ababa Ethiopia 2017
[3] T Esayas Exploration and Collection Characterization GeneticDiversity Analysis and Association of Traits for Yield and YieldComponents of Sugarcane (Saccharumspp) in Ethiopia [MasterThesis] Haramaya University Dire Dawa Ethiopia 2014
[4] J R Harlan Crops and Man American Society of AgronomyCrop Science Society of America Madison WI USA 2ndedition 1992
[5] R W Allard Principles of Plant Breeding JohnWilley and SonsNew York NY USA 2nd edition 1999
[6] B T Roach and J Daniels ldquoA review of the origin and improve-ment of sugarcanerdquo in Proceedings of Coper sugar InternationalSugarcane Workshop pp 1ndash32 1987
[7] P P Patil S S Patil and D U Patel ldquoGenetic diversity in sug-arcane (Saccharum spp Complex)rdquo Journal of Pharmacognosyand Phytochemistry vol 6 no 6 pp 570ndash573 2017
[8] R B Singh B Singh andR K Singh ldquoStudy of genetic diversityof sugarcane (Saccharum) species and commercial varietiesthrough TRAP molecular markersrdquo Indian Journal of PlantPhysiology vol 22 no 3 pp 332ndash338 2017
[9] R W Allard ldquoPopulation structure and sampling methodsrdquo inGenetic Resources in PlantsmdashTheir Exploration and Conserva-tion O H Frankel and E Bennett Eds IBP Handbook no 11pp 97ndash107 FA Davis Company Philadelphia PA USA 1970
[10] D R Marshall and A H D Brown ldquoOptimum samplingstrategies in genetic conservationrdquo inCrop Genetic Resources forToday and Tomorrow O H Frankel and J G Hawkes Eds pp53ndash80 Cambridge University Press 1975
[11] J G Hawkes Crop Genetic Resources Field Collection ManualPudoc Wagnningen Netherland 1980
[12] J G Hawkes The Diversity of Crop Plants Harvard UniversityPress London England 1983
[13] B P K Yerima Manual for Good Laboratory Practice Foodand Agriculture Organization of United Nation Addis AbabaEthiopia 1993
[14] A Walkley and I A Black ldquoAn examination of the degtjareffmethod for determining soil organic matter and a proposedmodification of the chromic acid titrationmethodrdquo Soil Sciencevol 37 no 1 pp 29ndash38 1934
[15] S R Olsen and L E Sommers ldquoPhosphorusrdquo in AgronomyMonograph A L Page et al Ed Methods of soil analysis part2 pp 403ndash430 ASA and SSSAMadisonWI USA 2nd edition1982
[16] R H Bray and L T Kurtz ldquoDetermination of total organic andavailable forms of phosphorus in soilsrdquo Soil Science vol 59 no1 pp 39ndash46 1945
[17] S Sahlemedhin andB Taye ldquoProcedures for soil and plant anal-ysisrdquo Tech Rep 74 National Soil Research Center EthiopianAgricultural Research Organization 2000
[18] F C Stickler S Weaden and AW Pauli ldquoLeaf area determina-tion in grain sorghumrdquo Journal of Agronomy and Crop Sciencevol 53 pp 187-188 1961
[19] Munsell Color ChartThe Munsell Book of Color Matte Collec-tion Munsell Color Chart NewWindsor NY USA 1990
[20] H O Hartley ldquoThe maximum F-ratio as a short cut test forheterogeneity of variancesrdquo Biometrika vol 37 pp 308ndash3121950
[21] L Pecetti P Annicchiarico and A B Damania ldquoBiodiversity ina germplasm collection of durumwheatrdquo Euphytica vol 60 no3 pp 229ndash238 1992
[22] L Pecetti and A B Damania ldquoGeographic variation intetraploid wheat (Triticum turgidum ssp turgidum convardurum) landraces from two provinces in Ethiopiardquo GeneticResources and Crop Evolution vol 43 no 5 pp 395ndash407 1996
[23] S K Jain C O Qualset G M Bhatt and K K Wu ldquoGeo-graphical patterns of phenotypic diversity in a world collectionof durumwheatsrdquoCrop Science vol 15 no 5 pp 700ndash704 1975
[24] K K Nkongolo L Chinthu M Malusi and Z Vokhiwa ldquoPar-ticipatory variety selection and characterization of sorghum(Sorghum bicolour (L) Moench) elite accessions from Malaw-ian gene pool using farmer and breeder knowledgerdquo AfricanJournal of Agricultural Research vol 3 pp 273ndash283 2008
[25] M B Semira ldquoInnovative methods and metrics for agricul-ture and nutrition actions soil quality and human nutritionin Ethiopiardquo Leverhulme Center for Integrative Research onAgriculture and Health (LCIRAH) 2017
[26] I A Khan A Khatri M A Siddiqui G S Nizamani and SRaza ldquoPerformance of promising sugarcane clone for yield andquality traits in different ecological zones of Sindhirdquo PakistanJournal of Botany vol 36 no 1 pp 83ndash92 2004
[27] M F Perera M E Arias D Costilla et al ldquoGenetic diversityassessment and genotype identification in sugarcane based onDNAmarkers andmorphological traitsrdquo Euphytica vol 185 no3 pp 491ndash510 2012
[28] C G Neil M Kay and C C Jack ldquoDiversity among mainlandUSA sugarcane cultivars examined by SSR genotypingrdquo Journalof the American Society of Sugar Cane Technologists vol 29 pp36ndash52 2009
[29] S SchenckMWCrepeauK KWu P HMooreQ Yu andRMing ldquoGenetic diversity and relationships in native HawaiianSaccharum officinarum sugarcanerdquo Journal of Heredity vol 95no 4 pp 327ndash331 2004
[30] R Gebretsadik H Shimelis M D Laing P Tongoona andN Mandefro ldquoA diagnostic appraisal of the sorghum farmingsystem and breeding priorities in Striga infested agro-ecologiesof Ethiopiardquo Agricultural Systems vol 123 pp 54ndash61 2014
[31] L E V Cesar A T Bruzi J A R Nunes et al ldquoGeneticdivergence of sugarcane varieties collected in the region ofLavras Minas Gerais Brazilrdquo Genetics and Molecular Researchvol 14 no 4 pp 14043ndash14049 2015
[32] C Grenier P J Bramel J A Dahlberg et al ldquoSorghums of theSudan analysis of regional diversity and distributionrdquo GeneticResources and Crop Evolution vol 51 no 5 pp 489ndash500 2004
[33] N V Nair R Balakrishnan and T V Sreenivasan ldquoVariabilityfor quantitative traits in exotic hybrid germplasm of sugarcanerdquoGenetic Resources and Crop Evolution vol 45 no 5 pp 459ndash464 1998
[34] R Nagarajan S Alarmelu and R M Shanthi ldquoStudies onvariation in interspecific hybrids of Saccharumrdquo Sugar Tech vol2 no 3 pp 42ndash46 2000
18 Advances in Agriculture
[35] D N Kamat and J R P Singh ldquoVariability in Sugarcane underrain fed conditionrdquo Sugar Technology Short Communicationsvol 3 no 1-2 pp 65ndash67 2001
[36] B S Rishipal A S M Chaudhary and S P Kadian ldquoStudieson character interrelationship in clonal generation of sugarcane(Saccharum complex hybrids)rdquo Indian Sugar vol 47 no 11 pp907ndash911 1998
[37] S Thippeswamy S T Kajjidoni P M Salimath and J V GoudldquoCorrelation and path analysis for cane yield juice quality andtheir component traits in sugarcanerdquo Short Communication vol5 no 1-2 pp 65ndash72 2003
[38] P S Verma S Pal and N K Karma ldquoGenetic variability andcorrelation studies in sugarcanerdquo Indian Sugar vol 49 pp 125ndash128 1999
[39] H A Esechie J W Maranville and W M Ross ldquoRelationshipof stalk morphology and chemical composition to lodgingresistance in sorghumrdquo Crop Science vol 17 no 4 p 609 1977
[40] P K Das B C Jena and N Nayak ldquoCharacter association andpath analysis of sugar yield in sugarcanerdquo Indian Sugar vol 46no 10 pp 805ndash808 1997
[41] K J Sreekumar M Kuriakose M Thomas D Alexander andS Santhakumari ldquoVariability heritability and correlation onstudies on the yield and quality characters of sugarcanerdquo IndianSugar vol 44 no 4 pp 243ndash249 1994
[42] J R Harlan J M de Wet and E G Price ldquoComparativeevolution of cerealsrdquo Evolution vol 27 no 2 pp 311ndash325 1973
[43] A Elings ldquoDurum wheat landraces from syria II patterns ofvariationrdquo Euphytica vol 54 no 3 pp 231ndash243 1991
[44] K Tato ldquoRainfall in Ethiopiardquo Ethiopian Geographical Journalvol 2 pp 28ndash36 1964
[45] A B L Stemler J R Harlan and J M J deWet ldquoThe sorghumsof ethiopiardquo Economic Botany vol 31 no 4 pp 446ndash460 1977
[46] P Annicchiarico L Pecetti and A B Damania ldquoRelationshipsbetween phenotypic variation and climatic factors at collectingsites in durum wheat landracesrdquo Hereditas vol 122 no 2 pp163ndash167 1995
[47] A B Damania L Pecetti C O Qualset and B O HumeidldquoDiversity and geographic distribution of adaptive traits inTriticum turgidum L (durum group) wheat landraces fromTurkeyrdquoGenetic Resources and Crop Evolution vol 43 no 5 pp409ndash422 1996
[48] A Ayana and E Bekele ldquoGeographical patterns of morpho-logical variation in sorghum (Sorghum bicolor (L) Moench)germplasm from Ethiopia and Eritrea quantitative charactersrdquoEuphytica vol 115 no 2 pp 91ndash104 2000
[49] D S Falconer Introduction to Quantitative Genetics JohnWilley and Sons Inc New York NY USA 3rd edition 1993
[50] R S Thorpe ldquoBiochemical analysis of geographical variationand racial affinitiesrdquo Biology Review vol 51 pp 407ndash452 1976
[51] M Farooq ldquoMorphological and agricultural characteristicsof sugarcane in relation to mechanized agriculture at NARCIslamabadrdquo Portu Rico Journal vol 1 pp 51ndash61 1989
[52] M Akhtar N N Elahi and M Ashraf ldquoEvaluation of exoticsugarcane varieties for agronomic characters and productivityrdquoPakistan Journal of Biological Sciences vol 4 no 1 pp 37ndash402001
[53] F R Piscitelli ldquoPrincipals caracteres exomorfologicos de cul-tures de cana de azucarrdquo TUC Iindustraly Agricos vol 7 pp 49ndash57 1994 (Spanish)
[54] A H D Brown and B S Weir ldquoMeasuring genetic variability inplant populationsrdquo in Isozyme in Plant Genetics Breeding S DTanksley and T J Orton Eds part A Elsevier Science 1983
[55] R Balakrishnan N V Nair and T V Sreenivasan ldquoA methodfor establishing a core collection of Saccharum officinarum Lgermplasm based on quantitative-morphological datardquo GeneticResources and Crop Evolution vol 47 no 1 pp 1ndash9 2000
[56] C A Nestor and B Q Reynaldo Sugarcane Variety Improve-ment in Southeast Asia and the Pacific for Enhanced andSustainable Productivity ndash Germplasm and Disease IndexingComponents Institute of Plant Breeding (IPB) College ofAgriculture (CA) UP Los Banos (UPLB) 2003
[57] A Maqbool J Muhammad and A Sagheer ldquoAgronomic traitsand morphological characteristics of some exotic varieties ofsugarcanerdquoPakistan Journal of Agricultural Research vol 19 no4 2006
after planting (STC10MAP) hand refractometer brix reading10 months after planting (HRBrix10MAP) millable stalkcount per hectare (MSCHA) single cane weight (SCW)number of internode (NOI) internode length (IL) stalkheight (SH) stalk diameter (SD) leaf length (LL) leaf width(LW) leaf area (LA) cane yield quintal per hectare (CYHA)and sugar yield quintal per hectare (SY) Data on 4 juicequality parameters ie brix percent (Brix) pol percent(pol ) purity percent (purity ) and sugar percent (SR) was also recorded For every accession ten plants wereused for recording data for quantitative characters whichwere recorded on plot basis Count data and cane yield wererecorded considering all cane stalks from the whole plot Forquantitative leaf characteristics measurement a proceduredeveloped in [18] was used
To categorize each accession morphologically sugarcanedescriptors adopted fromUSDA-ARSwere employed (GRIN2004) Data on 16 qualitative traits was recorded namelypresence or absence of bud cushion (BUDCUSHION) rel-ative degree of bud extension (BUDEXTEND) relative budshape (BUDSHAPE) relative shape of dewlap (DEWLAP-SHAP) type of outer auricle (AURICLEOUT) presence orabsence of stalk corky cracks (STALKCORKC) presenceor absence of stalk corky patches (STALKCORKP) relativeshape of ligule (LIGSHAPE) presence or absence of stalkgrowth cracks (STALKCRACK) presence or absence of budgroove (BUDGROOVE) relative plant erectness (ERECT)relative degree of internode alignment (INALIGN) relativeinternode shape (INSHAPE) colour of the leaves (LEAF-COLOR) colour of the exposed rind (RINDCOLE) andcanopy structure (CANOPY) Each accession was scored forthe most frequent character-state Leaf colour and colourof the exposed rind were examined and scored using theMunsell colour chart [19]
223 Statistical Analyses First analysis of variance wasmadefor the 21 characters for each location Homogeneity ofthe error variances among the locations was assessed in[20] F-max method for each of the 21 characters The testestablished the homogeneity of the error variances for allcharacters except TC4MAP For TC4MAP logarithmic datatransformation which is recommendable for continuousdata was used to homogenize the error variance Thendata of all the characters were subjected to test location-accession-related analysis of variance to determine effects oftest sites and genotype X environment (G X E) interactionThe results showed the nonsignificance of effects of testsites for SPC1MAP IL LW Pol sugar percent (SR) andthe nonsignificance of G X E interactions for SPC2MAPSCW Brix Pol and Brix10MAP For the other charactersboth test site effects and G X E interactions were significant(Plt005)
Analysis of variance was made for 21 quantitative char-acters following the procedure used in [21 22] The meansquares of the regions were tested against pooled meansquares of accessions within regions The pooled meanssquares for accessions within regions of origin and the meansquares of accessions within each region were tested against
the pooled within-region error mean squares Means rangesfor means and percent coefficients of variation for all thecharacters were computed for each region of origin and forthe entire data The regional means were compared usingDuncanrsquos multiple range testing Correlations between thecharacters were computed at three levels First correlationsof the characters were assessed based on the 211 accessionmeans Then interregion correlation was computed usingthe means of characters for each region Finally a series ofintraregion correlation coefficient matrices were obtained foreach region using the accession means from that region forthe characters
For qualitative characters phenotypic frequency distri-butions were worked out for all the sample germplasm andlocations The ShannonndashWeaver diversity index (H1015840) wascomputed using the phenotypic frequencies to assess thephenotypic diversity for each character for all accessions TheShannonndashWeaver diversity index as described in [23] is givenas
1198671015840 = minus119899
sum119894=1
119901119894log119890(119901119894) (1)
where pi is the proportion of accessions in the 119894th class of ann-class character and n is the number of phenotypic classesof traits Each 1198671015840 value was divided by its maximum value(logen) and normalized in order to keep the values between0 and 1 By pooling various characters across the regions theadditive properties of 1198671015840 were used to evaluate diversity ofregions and characters within the population The averagediversity index (H1015840) over n traits was estimated as H1015840 =sumH1015840n
3 Results and Discussion
31 Collection of Sugarcane Germplasm In Ethiopia thehistory of sugarcane cultivation by smallholder farmers pre-ceded that of commercial cultivation As documented in thehistory of a monastery in Northern Ethiopia sugarcane hasbeen grown in the country since the 16th century [2] In thisstudy local sugarcane germplasm exploration and collectionwere conducted all across Ethiopia in the regional statesMore than 300 local sugarcane genotypes were collectedduring 20102011 and passport data of the genotypes ispresented in Supplemental Table 1 The collected germplasmwere planted at five locations across the country with respectto their collection area These are Wondogenet AgriculturalResearchCenter JimmaAgricultural ResearchCenterMechaWereda Agricultural Bureau nursery field at Picolo AbaySirinka Agricultural Research Center (at Kobo Subcenter)and research field of Haramaya University at Diredawa(Tonny Farm) The clones have been monitored during the201112 season for symptoms of major diseases and insectsThese clones were transferred to commercial sugarcane plan-tation estates at Wonji and Metehara for further selectionand maintenance No major diseases and insect pests weredetected in all the germplasm collected except sometimestermites and borer were observed (Supplemental Table 2)
Advances in Agriculture 5
Table 2 Mean soil physical and chemical properties of sugarcane fields in the surveyed regions
Regions Soil chemical properties Texture ()PH EC(dsm) OC N P (ppm) K(ppm) Sand Silt Clay
Most of the sampled germplasm had acceptable levels of juicerefractometer readings (expressed in degree brix) relative tothe history of the ages of the samples (Supplemental Table 1)The number of harvestablesaleable cane stalks obtained inplant cane and ratoon crops and the number of harvestableratoon crops showed variation region to region and zone tozone and sometimes district (ldquoweredardquo)-wise There is alsovariation in maturity time of the clones both in plant caneand in ratoon crops (Supplemental Table 2)These are mainlydepending on the variety soil condition cultural practicesand climatic conditions
The genotypes collected from the respective zones inthe study regions were recorded by their local namesSometimes similar varieties bear different names at differentlocalities and vice versa Farmers mentioned a broad rangeof sugarcane landraces that had been grown in the areasand maintained for generations Some of the landraces werecommonly recognized by most farmers within and acrosszones of the respective regions whereas some were rarevarieties known only by few farmers The large number oflandraces observed during the current study demonstratesthe existence of diverse genetic resources of sugarcane inEthiopia The diversity could have evolved presumably due todiverse climatic conditions low input management systemshigh pests and disease pressure and continuous selectionby farmers Smallholder farmers face diverse environmentstresses and have multiple production objectives that affectselection of genotypes [24] There were no formally releasedimproved sugarcane varieties grown by farmers in the studyregions except in few places This was mainly observedin SNNPR at Wolayta and Kembata-Tembaro zones Thesevarieties have been informally introduced to these areas byseasonal workers employed at the sugar estates especially atWonji and Metehara
The collected landraces would serve to broaden thegenetic base of the available sugarcane germplasm Thesugar industry in Ethiopia is currently establishing sugarcanebreeding programTherefore collection and efficient charac-terization of germplasm are vital to strengthen the breedingprogram
311 Physical and Chemical Properties of Soils of Sugar-cane Germplasm Collection Areas Table 2 summarizes thephysical and chemical properties of soils in the sugarcane
production fields of surveyed areas Sugarcane productionby smallholder farmers in the surveyed areas is conductedacross the seasons where harvesting and planting are done aslong as there is available moisture Therefore the soil samplesrepresent the sugarcane production season and the altitudinalranges of the respective areas
Generally the organic carbon content of soils in sugarcaneproducing areas across the country is low to very low (2-4lt2) Amhara Gambella SNNP Somali and Tigray Regionsare in a very low category (lt2) Similarly across regions thetotal N content is low except Harari and Oromia regionswhere the average total N is medium This problem shouldbe addressed either with augmenting the soil with inorganicfertilizers or through appropriate management practices likecrop residue management Available phosphorus is high inAmhara Gambella and SNNP Regions whereas moderatevalues were observed in Harari Oromia and Tigray RegionsGenerally the level of phosphorus in soils of sugarcaneproducing areas in these regions is adequate In Benshangul-Gumz and Somali Regions low level of phosphorus wasrecorded which should be addressed with due attentionAvailable K is sufficient for normal sugarcane growth in soilsof all regions Relatively lower values of K were recorded insoils of Harari and Tigray Regions It should be noted thatorganic carbon EC PH and variability of P andK are affectedby rainfall and temperature that are seasonal and the valuesrecorded refer to the seasonwhen the soil samples were takenGenerally Ethiopian soils are deficient in various essentialnutrients like boron nitrogen phosphorus potassium sulfurzinc and copper although severity differs from region toregion [25] Particularly the loss of P and N resulting fromthe use of dung and crop residues for fuel is demonstrated tobe equivalent to the total amount of commercial fertilizer use
Across regions in Ethiopia generally clay type of soildominates followed by silt except in Gambella Region wherethe dominant soil texture was sand Soil management optionsrelated to clay soil may address soil related problems insmallholder sugarcane growing areas across Ethiopia
32 Diversity of Sugarcane Landraces
321 Quantitative Characters
Univariate Statistics Analysis of variance depicted highlysignificant differences (p lt 001) between accessions pooled
6 Advances in Agriculture
over the regions for the 21 characters of the 211 sugarcaneaccessions studied and significant difference for the 6 regionsof origin for 6 characters (Table 3) The results suggestedthe occurrence of significant phenotypic variation betweenthe accessions as a whole Significant variations of differentsugarcane stalk and juice quality characters was also reportedin similar studies conducted elsewhere [26 27] Region-wise partitioning of the variance indicated significant within-region differences (p lt 005) among the populations withinalmost all regions for the characters NOI IL SH SD LLLW LA Brix Pol Purity SR MSCHA SCW CYHAand SY for 8 characters within Tigray for TC4MAP andTC5MAP within Oromiya and SNNPR Brix10MAP showedno significant difference within all regions
In general within-region variation was greater for stalkdiameter single cane weight millable stalk count and caneand sugar yield than for other characters for all the regionsAssuming that a significant portion of the phenotypic vari-ation is genetic it would be possible to make selection forany of this group of characters within a particular regionIt was apparent that the variance between accessions pooledover regions was greater than between regions Therefore inorder to sample the variation effectively it would be necessaryto sample the variable populations from different localitiesin a region The study in [28] on genetic diversity amongmain land USA sugarcane cultivars showed high geneticdiversitywithin populations of different regions than betweenpopulations
Duncanrsquos multiple range testing for regional means for allthe characters is shown in Table 4 Relatively much differenti-ationwas observed for stemdiameter and juice purity percentcompared to other parameters Important yield componentslike MSCHA SCW SH and SD have shown higher valuesin Tigray SNNPR and Oromiya regions High cane andsugar yield were recorded for accessions in Tigray regionfollowed by SNNPR Oromiya and introduction which werestatistically at par Statistically similar lower values of caneand sugar yield were recorded in Amhara and Benshangulcollections
Higher value of sucrose percent in cane was also observedfor accessions from Tigray and the lowest value was inAmhara accessions while other regions were statisticallysimilar Generally for most of the characters high valuewas obtained from accessions in Tigray region followed bySNNPROromiya and introduction Therefore the materialsfrom these regions can be used for selection of cane and sugaryield per se and in breeding program The local accessions inmany of the regions showed superiority over the introducedcommercial varieties currently in production for most of thecharacters including cane and sugar yield (Table 4)This maybe due to the adaptation and coevolution of the materialswith different stresses for long period of time in the countryrsquosagroecology These locally adapted genes conferring yieldadvantage could be harnessed through strategic crossing andselection for improvement of the crop Superiority of locallandraces over introduced varieties was reported [29 30]
The range of variation of the accession means establishedwide variation between the regions and the accessions withinthe regions for the characters studied (Table 5) Accordingly
the maximum score was 41 times the minimum for sproutcount one month after planting 32 for sprout count twomonths after planting 25 for tiller count four months afterplanting 22 for sugar yield 20 for cane yield 12 for millablestalk count 8 for stalk count 10 months after planting 7for single cane weight 6 for tiller count five months afterplanting and 2 for other yield components and sugar qual-ity parameters except hand refractometer brix reading tenmonths after planting and juice purity percent which scored1 The same trend was observed between accessions withina particular region However accessions within AmharaBenshangul-Gumz and introduction showed wider ranges ofvariation than those accessions within other regions for themajority of the characters The wide range of geographic andclimatic features of the regions in Ethiopia and the originaldifferentiation and further farmers selection for productionniches and uses for hundreds of years may have resultedin the accessions to possess a tremendously high degree ofmorphological variation The reason for introduced varietiesshowing high variation also could be due to broader geo-graphic spectrum fromwhere theywere initially acquired anddifference in their parental source and genealogical historyIt was demonstrated even with these limited numbers ofsamples (15 genotypes) that introduction of genetic materialshas practically broadened the genetic base of the sugarcanegermplasm for the breeding program
Coefficient of variation measured as the ratio of standarddeviation to the corresponding overall mean and expressedas percentage is useful to compare different charactersmeasured in different units It is also useful to compare samecharacter in different groups of populations with differentsample size mean and variance or different characters indifferent populations In the current study high coefficientsof variation were observed between regions and within eachregion for sprout as well as tiller counts in the monthsthe data was recorded stalk count 10 months after plant-ing and millable stalk count at harvest cane and sugaryield single cane weight and leaf area (Table 6) Thoughaccessions from a particular region were more variable fora specific character compared to other regions accessionsfromAmhara Benshangul-Gumz introduction and SNNPRwere more variable than accessions from other regions Thisresult demonstrated the tremendous variability of sugar-cane germplasm from these regions The accessions fromTigray and Oromiya had low coefficients of variation formany characters indicating relatively high within-regionuniformity The high coefficients of variation observed formost of the characters agreed well with those reportedin [31] for indigenous sugarcane of Brazil Similar resultswere also reported in sorghum [32] The different levels ofregional variability of a particular character could be due todifferences in forces of selection andor differences in theintensity of a particular selecting force It could also be dueto random effect and reduced sample parents of parentalmaterials moved differently by various human populationmigration and the interaction of these with Darwinianforces
Results of this study suggested the presence of wide rangeof variations of characters determined by univariate statistics
This is in agreement with previous studies on sugarcanegermplasm [33ndash35]
This is the first study to report information on geo-graphical pattern of variation in Ethiopian local sugarcanegermplasm which has been lacking so far Our results pro-vided experimental evidence on occurrence of geographicalvariation and significant within-region variation where it washigh in the regions such as Amhara Benshangul-Gumz andSNNPR
The overall patterns of similarity or difference betweenregions seemed to depend on environmental factors such asrainfall temperature length of growing season and altitudeWide variability of millable stalk count at harvest singlecane weight and plant height was observed among regionswhere these characters demonstrated high correlation withcane and sugar yield (Table 7) Therefore breeders can utilizeaccessions of regions showing variability for these charactersin selection programs and to design breeding strategies toproduce varieties with best commercial merits
Bivariate Statistics Correlation coefficients worked out onthe 21 quantitative characters are shown for the entire data(Table 7) between regions (Table 8) and within regions(data not shown) Results of the association studies for theentire data showed that cane and sugar yield had highlysignificant positive correlation with most of the quantitativestalk characters and juice quality parameters However theassociation of tiller counts 4 and 5 months after plantingstalk count 10 months after planting millable stalk countat harvest single cane weight and plant height with caneand sugar yield was stronger suggesting these componentcharacters asmain contributing factor to cane and sugar yieldIn fact sugar yield is the product of cane yield and sucrose per-cent cane surely being influenced with components affectingcane yield Similar trend was reported by earlier sugarcaneworkers [36 37] In sugarcane the cane and sugar yields areconsidered to be the complex characters The information onthe phenotypic interrelationship of cane yield and sugar yieldwith their component character per se would be of immensehelp to sugarcane breeders
Stem diameter had significant positive association withboth cane and sugar yield indicating the significance ofthis trait in improving both cane and sugar yields Similarreports have beenmade by earlier sugarcane workers [37 38]Stalk diameter was also positively and significantly correlatedwith plant height It is one of the traits that are related tolodging resistance [39] and the positive associations withheight would help in reducing the chance of lodging as heightincreases Among the physiological attributes included in thestudy leaf area had highly significant positive associationwith cane yield Similar observation was also made in [37]Sucrose percent which is an important juice quality attributehad highly significant positive association with sugar yieldand simultaneously cane yield also had highly significantassociation with sugar yield Based on the magnitude ofcorrelation coefficient values it can be said that cane yield ismuch more important than sucrose percent in determiningthe sugar yield and the same has also been reported by othersugarcane workers [37 40] Similar to observations made in
[37 41] results of the current study showed that all the qualityparameters like sucrose percent pol percent brix percentand purity percent were significantly correlated in positivedirection This indicated any of these juice quality traitscould be considered for selection leading to the simultaneousimprovement in the remaining quality traits
Ten of the 21 characters also showed significant positivecorrelations with altitude of the collection sites (Table 7) Asdescribed in [42] ecological characteristics have influencedthe genotypic constitution of landraces during domesticationand hence a relationship exists between the agroecology inthe collection site and the morphological characteristics ofthe landraces Thus positive correlation between collectionsite variables and plant characteristics would imply that thevariation between accessionsmay be related to agroecologicalvariations among the collection sites [43]
The correlation coefficient between number of internodesand altitude was negative and significant indicating thatother environmental factors (other than altitude) andornonenvironmental factorsmight account for the variation forthis particular character Since temperature decreases withan increase of altitude in Ethiopia [44 45] it is more likelythat temperature has exerted strong selection pressure innumber of internodes in the genotypes Information on therelationships between environmental factors of the collect-ing sites and morphophysiological variation of germplasmcould enhance the understanding of evolutionary adaptivepatterns which could assist germplasm collectors and users[46]
Character associations between region and within regionalso followed similar fashion like that of the entire dataNumber of sprout tiller and stalk counts and sugar qualityparameters showed significant positive correlations with caneand sugar yield
Though not significant and strong the correlation of leaflength and width and leaf area (leaf area = leaf length Xleaf width X 075) with cane and sugar yield was negativein the interregion correlations This kind of relationship wasalso exhibited between these leaf characteristics and yieldcomponents like tiller and stalks counts plant height andsingle cane weight and stalk diameter and sugar qualityparameters This may have been attributed to the variationof temperature with altitude and other environmental factorsacross the regions resulting in such type of relationshipSuch relationship also calls for the need to have enough dataon environmental variables of the collection sites such astemperature and rainfall if germplasm collection is requiredtomeet its objectives effectively and efficiently Unfortunatelydata on such useful environmental variables like temperatureand rainfall are lacking in the passport data of the currentEthiopian sugarcane germplasm collections The significanceof environmental factors of the collection sites as veryimportant determinants in structuring morphophysiolog-ical variations was reported in tetraploid wheat [22 4748]
Association of characters among yield its componentsand other economic traits is important for the interpretationof the patterns of variation and making selection in breed-ing program and combining several desirable attributes It
suggests the advantage of a scheme of selection for morethan one character at a time The correlation between char-acters may arise from linkage or from developmental geneticinteractions with orwithout a purely phenotypic componentIt could arise also due to differential phenotypic plasticityof characters themselves Within the limit of experimentalerror and environmental effects high correlation coefficientsbetween characters may show that the characters sharesome common element of genetic control (ie pleiotropylinkage) between genes or else from independently controlledcharacters responding similarly to geographic variation inselection pressures [49] The interregion correlation coef-ficient between the characters measures the consistency oftheir patterns of regional variation while the intraregioncorrelation coefficient measures the association arising fromgenetic factors but is not affected by regional variation[50] Since this study showed significant positive correlationsintraregionally for some character combinations it wouldseem that common genetic control might be playing a role inbringing about correlations between the various charactersIt appeared that different response to regional variationwas playing a greater role than different genetic controlas shown by the many more significant and moderate tohigh correlation coefficients intraregionally than interregion-ally
322 Qualitative Characters The frequency distribution forthe 16 qualitative characters of germplasm samples by regionsin Ethiopia and of the exotic accessions is shown in Supple-mental Table 5 For the purpose of avoiding redundancy andimproving readability the discrete characters are presented bygrouping related traits together
BudCushion andRelativeDegree of BudExtension Evaluationof the presence and absence of bud cushion revealed thatgreat proportion (70) of sugarcane landraces from Ethiopiado not have bud cushion Majority of the sample germplasmin all regions belong to this phenotypic class Approximatelyone-half (53) of the collection have their bud extendedabove the growth ring followed by those (41) having budtouching the growth ring
Relative Bud Shape Among the phenotypic classes for thischaracter tall deltoid was dominant (31) followed by ovate(17) and narrow ovate (12) While local accessions inmost regions had tall deltoid bud shape 50 of the intro-duced accessions belong to two phenotypic classes roundwith central germpore (30) and 20 having pentagonalbud shape Equal proportions (19) of the samples fromBenshangul-Gumz were Ovate and Ovate with emarginatebasal wing
Canopy Structure Relative Plant Erectness Colour of theLeaves and Colour of the Exposed Rind Out of the eightphenotypic classes observed compact tip droopy was themost frequent (35) Open semidroopy and open droopyclasses had fairly equal distribution With regard to relativeplant erectness seventy-five percent of the germplasm werefound to fall in twophenotypic classes erect (41) and almost
erect (34) for the whole region and country Most of theaccessions (90) showed light green or green leaf colourbut 20 from introduced collections were greenish yellowAmong phenotypic classes for colour of the exposed rindbrownish yellow yellow brown and yellowish green were themost frequent with fairly equal distribution
Relative Shape of Dewlap Double crescent had larger fre-quency (22) while squarish descending and flaring typeof dewlap shape had fairly equal proportions Seventeenpercent of germplasm from Tigray region had squarishdeltoid dewlap shape
Relative Degree of Internode Alignment and Internode ShapeForty-four percent of the sample germplasm were slightlyzigzag while there was almost equal frequency for straightand zigzag internode alignment for the entire region andcountry Straight or nearly straight (slightly zigzag) internodealignment of cane stalks is a very important character formechanized farming and postharvest handling in sugarcane[51] Forty percent of the collections were observed havingconcave-convex type of internode shape while 29 wascylindrical and 14 conoidal Other classes had fairly equalproportion
Type of Auricle and Relative Shape of Ligule An equalproportion of the germplasm totally constituting 84 ofthe collection belong to four phenotypic classes namelythose with no auricle and others with transitional and shortand lanceolate auricles Sixteen percent of the collectionsalso showed equal frequency for the remaining classesBenshangul-Gumz and Tigray regions had each equal 13accessions with falcate type of auricle Sixty-seven percentof the collections belong to three ligule shape classes cres-cent with lozenge (27) broad-crescent (21) and linear-crescent (19) The germplasm had almost equal proportionfor the other phenotypic classes of this character Nineteenpercent of the accessions fromBenshangul-Gumz region haddeltoid shape of ligule
Stalk Corky Cracks Corky Patches Growth Cracks and BudGroove Eighty-nine percent of the germplasm across regionshad no stalk corky cracks with only eleven percent exhibitingthis character All the germplasm in Gambella Harari andSomali had no stalk corky cracks On the other hand eighty-one percent of the germplasm for the whole region showedthe presence of stalk corky patches All the local germplasmin Benshangul-Gumz Gambella and Tigray had stalk corkypatches whereas 75 of the germplasm from Harari Region67 from Somali Region and 24 from Oromia Regionhad no stalk corky patches In the entire region 75 ofthe germplasm had no stalk growth cracks with only 25showing this character Sixty-three percent of the germplasmwere found to have bud groove Howevermost of the samplesin Harari (75) Amhara (63) and Oromia (53) Regionshad no bud groove
The results of this study indicated the wide distributionof phenotypic classes for the characters considered whichindicates the existence of different races and combination
Advances in Agriculture 15
of races in local genotypes of different regions in EthiopiaThis diversity of germplasmbased onphenotypicmarkers canhelp sugarcane breeders in identifying the identity of geno-types as well as maintaining genetic diversity Differencesin morphological characters of different sugarcane varietieshave been reported [52] Piscitelli [53] showed importantexomorphological qualitative characters of sugarcane varietythat are not influenced by environmental factors and thus canbe used as a selection tool in any breeding program
The amount of phenotypic diversity estimates based onthe ShannonndashWeaver diversity index (H1015840) are shown inTable 9 The 16 characters differed in their distribution aswell as the amount of variation Individual traits showed adifferent pattern of variation among accessions Estimatesof H1015840 for individual characters varied from 049 for stalkcorky cracks to 100 for bud groove with an overall meanof 080 Most of the characters were highly polymorphicwhereas relative plant erectness and presence or absenceof stalk corky patches scored moderate values The highdiversity values for the characters showed a wide variabilityamong genotypes According to [54] this index is also usedin genetic resource studies as a convenient measure of bothallelic richness and allelic evenness when using geneticdata
The highest diversity computed was on presence orabsence of bud groove with a diversity index of 100indicating that this character is very variable in the localsugarcane germplasm On the other hand the descriptorcomputed with the lowest diversity index value (049) wasthe presence or absence of stalk corky cracks indicatingthat this character is selected in the development of thegenotypes In agreement with the present study [55] workingon S officinarum accessions from the world collection ofsugarcane germplasm obtained a low index of 025 and 035for presence or absence of stalk corky patches and cracksrespectively
Their report also showed high diversity index values forinternode alignment growth cracks internode shape budcushion bud extension bud groove and ligule shape Sim-ilarly [56] experimenting on field collections of sugarcaneaccessions in Philippine reported high ShannonndashWeaverdiversity for stalk growth cracks bud cushion stalk corkypatch stalk corky cracks bud extension and auricle outershape Unlike the present study they have reported high indexfor stalk corky cracksMedium index values were reported forstalk internode shape and low index values for ligule shapeand bud shape High variability of sugarcane varieties fordewlap shape was also reported in [57]
The H1015840 pooled across characters by region ranged from079 in Oromiya to 085 in Tigray with an overall aver-age of 081 (Table 8) Most of the regions showed highShannonndashWeaver diversity index The populations that hadthe highest H1015840 were that of Tigray and introduction The H1015840averaged over regions for different characters was found torange from 044 for presence or absence of stalk corky cracksto 094 for relative degree of internode alignment and budgroove with an overall average of 081 (Table 9)
Assessment of the ShannonndashWeaver index also showedsensitivity to both the number of descriptor classes and the
distribution within phenotypic classes in each region Amongregions the mean values for the ShannonndashWeaver index arenot significantly different but differences were found whenindividual characters were considered These observationssuggest that some regions have high diversity for particulartraits while in other regions selective pressures might havereduced the variation to certain genotypes
4 Conclusions
The large number of landraces observed during the cur-rent study demonstrates the existence of diverse geneticresources of sugarcane in Ethiopia The landraces wouldserve to broaden the genetic base of the available sugarcanegermplasm Generally distribution of the overall phenotypicdiversity among sugarcane landraces in Ethiopia is unevenOur results showed that there was a wide range of variationthat existed in the local sugarcane landraces studied both atregional and within region levels Selection and differenti-ation of types might have taken place along a geographicalpattern perhaps associated with climate and use The presentstudy contributes to updating sugarcane descriptors adoptedfrom USDA-ARS as well as Bioversity passport data forthe future collection and evaluation As Ethiopian SugarCorporation is in the course of launching breeding programthe observed substantial variation of landraces would enablesugarcane breeders to design and practice breeding andselection programs to improve the crop
Data Availability
The data used to support the findings of this study areavailable from the corresponding author upon request
Conflicts of Interest
The authors declare that they have no conflicts of interest
Acknowledgments
The authors are grateful for the financial grant of SugarCorporation of Ethiopia Many thanks are due to EthiopianBiodiversity Institute (EBI) for its technical advice duringcollection of the local sugarcane genotypes Sincere thanksare forwarded to Ethiopian smallholder farmers who pre-served sugarcane germplasm and shared their knowledgeand experience Regional Zonal and District (ldquoWeredardquo)Agriculture Bureaus across Ethiopia are acknowledged fortheir support during the study
Supplementary Materials
These supplementary materials contain different tables sup-porting the results of the study These include passport dataof sugarcane germplasm collected and related informationon the materials local sugarcane germplasm collected andused formorphological diversity study introduced sugarcane
16 Advances in Agriculture
Table9Sh
anno
n-WeaverD
iversityIndices(H1015840)a
ndmeanandsta
ndarderror(plusmnS
E)of
land
racesineach
region
basedon
16discretecharacters
Characters
Amha
raBe
nsha
ngul-G
umz
Introd
uced
Oromia
SNNP
Tigray
ELb
Total
Presence
orabsenceo
fbud
cushion(BUDCUSH
ION)
083
095
095
089
082
098
090
088
Relativ
edegreeo
fbud
extension(BUDEX
TEND)
077
079
093
074
072
087
080
078
Relativ
ebud
Shape(BU
DSH
APE
)078
096
089
081
085
078
084
078
Canop
ystructure(C
ANOPY
)089
098
095
081
078
096
089
084
Relativ
eshape
ofdewlap(D
EWLA
PSHAP)
087
094
092
090
082
096
090
086
Relativ
eplant
erectness(ER
ECT)
080
082
050
067
071
088
073
064
Relativ
edegreeo
finterno
dealignm
ent(IN
ALIGN)
096
076
098
097
098
100
094
098
Relativ
einterno
deshape(IN
SHAPE
)086
069
095
077
083
091
083
082
Color
oftheleaves(LE
AFC
OLO
R)087
071
090
064
074
071
076
076
Type
ofou
tera
uricle(A
URI
CLE
OUT)
080
092
087
079
083
093
086
082
Relativ
eshape
ofligule(LIGSH
APE
)086
094
082
085
090
083
087
083
Presence
orabsenceo
fstalkcorkyc
racks(ST
ALK
CORK
C)
044
054
035
055
051
025
044
049
Presence
orabsenceo
fstalkcorkyp
atches(STA
LKCORK
P)066
000
072
079
059
087
060
069
Presence
orabsenceo
fstalkgrow
thcracks
(STA
LKCRA
CK)
066
100
088
062
088
099
084
080
Color
ofthee
xposed
rind(R
INDCOLE
)085
096
089
084
086
076
086
085
Presence
orabsenceo
fbud
groo
ve(BUDGRO
OVE)
095
100
072
100
100
099
094
100
Means
080plusmn0
03
081plusmn0
06
083plusmn0
04
079plusmn0
03
080plusmn0
03
085plusmn0
05
081plusmn0
03
080
b EL=
averageo
fentire
locatio
n
Advances in Agriculture 17
varieties used for morphological diversity study and fre-quency distribution for 16 qualitative characters in sugarcaneby regions in Ethiopia and country of sources of introducedgenotypes (Supplementary Materials)
References
[1] ldquoSpreading the word about clean solutions from sugarcanrdquoAccessed 04072018 httpwwwsugarcaneorg
[2] Central Statistics Agency (CSA) ldquoAgricultural sample surveyArea and production of major crops private peasant holdingsrdquoStatistical Bulletin Addis Ababa Ethiopia 2017
[3] T Esayas Exploration and Collection Characterization GeneticDiversity Analysis and Association of Traits for Yield and YieldComponents of Sugarcane (Saccharumspp) in Ethiopia [MasterThesis] Haramaya University Dire Dawa Ethiopia 2014
[4] J R Harlan Crops and Man American Society of AgronomyCrop Science Society of America Madison WI USA 2ndedition 1992
[5] R W Allard Principles of Plant Breeding JohnWilley and SonsNew York NY USA 2nd edition 1999
[6] B T Roach and J Daniels ldquoA review of the origin and improve-ment of sugarcanerdquo in Proceedings of Coper sugar InternationalSugarcane Workshop pp 1ndash32 1987
[7] P P Patil S S Patil and D U Patel ldquoGenetic diversity in sug-arcane (Saccharum spp Complex)rdquo Journal of Pharmacognosyand Phytochemistry vol 6 no 6 pp 570ndash573 2017
[8] R B Singh B Singh andR K Singh ldquoStudy of genetic diversityof sugarcane (Saccharum) species and commercial varietiesthrough TRAP molecular markersrdquo Indian Journal of PlantPhysiology vol 22 no 3 pp 332ndash338 2017
[9] R W Allard ldquoPopulation structure and sampling methodsrdquo inGenetic Resources in PlantsmdashTheir Exploration and Conserva-tion O H Frankel and E Bennett Eds IBP Handbook no 11pp 97ndash107 FA Davis Company Philadelphia PA USA 1970
[10] D R Marshall and A H D Brown ldquoOptimum samplingstrategies in genetic conservationrdquo inCrop Genetic Resources forToday and Tomorrow O H Frankel and J G Hawkes Eds pp53ndash80 Cambridge University Press 1975
[11] J G Hawkes Crop Genetic Resources Field Collection ManualPudoc Wagnningen Netherland 1980
[12] J G Hawkes The Diversity of Crop Plants Harvard UniversityPress London England 1983
[13] B P K Yerima Manual for Good Laboratory Practice Foodand Agriculture Organization of United Nation Addis AbabaEthiopia 1993
[14] A Walkley and I A Black ldquoAn examination of the degtjareffmethod for determining soil organic matter and a proposedmodification of the chromic acid titrationmethodrdquo Soil Sciencevol 37 no 1 pp 29ndash38 1934
[15] S R Olsen and L E Sommers ldquoPhosphorusrdquo in AgronomyMonograph A L Page et al Ed Methods of soil analysis part2 pp 403ndash430 ASA and SSSAMadisonWI USA 2nd edition1982
[16] R H Bray and L T Kurtz ldquoDetermination of total organic andavailable forms of phosphorus in soilsrdquo Soil Science vol 59 no1 pp 39ndash46 1945
[17] S Sahlemedhin andB Taye ldquoProcedures for soil and plant anal-ysisrdquo Tech Rep 74 National Soil Research Center EthiopianAgricultural Research Organization 2000
[18] F C Stickler S Weaden and AW Pauli ldquoLeaf area determina-tion in grain sorghumrdquo Journal of Agronomy and Crop Sciencevol 53 pp 187-188 1961
[19] Munsell Color ChartThe Munsell Book of Color Matte Collec-tion Munsell Color Chart NewWindsor NY USA 1990
[20] H O Hartley ldquoThe maximum F-ratio as a short cut test forheterogeneity of variancesrdquo Biometrika vol 37 pp 308ndash3121950
[21] L Pecetti P Annicchiarico and A B Damania ldquoBiodiversity ina germplasm collection of durumwheatrdquo Euphytica vol 60 no3 pp 229ndash238 1992
[22] L Pecetti and A B Damania ldquoGeographic variation intetraploid wheat (Triticum turgidum ssp turgidum convardurum) landraces from two provinces in Ethiopiardquo GeneticResources and Crop Evolution vol 43 no 5 pp 395ndash407 1996
[23] S K Jain C O Qualset G M Bhatt and K K Wu ldquoGeo-graphical patterns of phenotypic diversity in a world collectionof durumwheatsrdquoCrop Science vol 15 no 5 pp 700ndash704 1975
[24] K K Nkongolo L Chinthu M Malusi and Z Vokhiwa ldquoPar-ticipatory variety selection and characterization of sorghum(Sorghum bicolour (L) Moench) elite accessions from Malaw-ian gene pool using farmer and breeder knowledgerdquo AfricanJournal of Agricultural Research vol 3 pp 273ndash283 2008
[25] M B Semira ldquoInnovative methods and metrics for agricul-ture and nutrition actions soil quality and human nutritionin Ethiopiardquo Leverhulme Center for Integrative Research onAgriculture and Health (LCIRAH) 2017
[26] I A Khan A Khatri M A Siddiqui G S Nizamani and SRaza ldquoPerformance of promising sugarcane clone for yield andquality traits in different ecological zones of Sindhirdquo PakistanJournal of Botany vol 36 no 1 pp 83ndash92 2004
[27] M F Perera M E Arias D Costilla et al ldquoGenetic diversityassessment and genotype identification in sugarcane based onDNAmarkers andmorphological traitsrdquo Euphytica vol 185 no3 pp 491ndash510 2012
[28] C G Neil M Kay and C C Jack ldquoDiversity among mainlandUSA sugarcane cultivars examined by SSR genotypingrdquo Journalof the American Society of Sugar Cane Technologists vol 29 pp36ndash52 2009
[29] S SchenckMWCrepeauK KWu P HMooreQ Yu andRMing ldquoGenetic diversity and relationships in native HawaiianSaccharum officinarum sugarcanerdquo Journal of Heredity vol 95no 4 pp 327ndash331 2004
[30] R Gebretsadik H Shimelis M D Laing P Tongoona andN Mandefro ldquoA diagnostic appraisal of the sorghum farmingsystem and breeding priorities in Striga infested agro-ecologiesof Ethiopiardquo Agricultural Systems vol 123 pp 54ndash61 2014
[31] L E V Cesar A T Bruzi J A R Nunes et al ldquoGeneticdivergence of sugarcane varieties collected in the region ofLavras Minas Gerais Brazilrdquo Genetics and Molecular Researchvol 14 no 4 pp 14043ndash14049 2015
[32] C Grenier P J Bramel J A Dahlberg et al ldquoSorghums of theSudan analysis of regional diversity and distributionrdquo GeneticResources and Crop Evolution vol 51 no 5 pp 489ndash500 2004
[33] N V Nair R Balakrishnan and T V Sreenivasan ldquoVariabilityfor quantitative traits in exotic hybrid germplasm of sugarcanerdquoGenetic Resources and Crop Evolution vol 45 no 5 pp 459ndash464 1998
[34] R Nagarajan S Alarmelu and R M Shanthi ldquoStudies onvariation in interspecific hybrids of Saccharumrdquo Sugar Tech vol2 no 3 pp 42ndash46 2000
18 Advances in Agriculture
[35] D N Kamat and J R P Singh ldquoVariability in Sugarcane underrain fed conditionrdquo Sugar Technology Short Communicationsvol 3 no 1-2 pp 65ndash67 2001
[36] B S Rishipal A S M Chaudhary and S P Kadian ldquoStudieson character interrelationship in clonal generation of sugarcane(Saccharum complex hybrids)rdquo Indian Sugar vol 47 no 11 pp907ndash911 1998
[37] S Thippeswamy S T Kajjidoni P M Salimath and J V GoudldquoCorrelation and path analysis for cane yield juice quality andtheir component traits in sugarcanerdquo Short Communication vol5 no 1-2 pp 65ndash72 2003
[38] P S Verma S Pal and N K Karma ldquoGenetic variability andcorrelation studies in sugarcanerdquo Indian Sugar vol 49 pp 125ndash128 1999
[39] H A Esechie J W Maranville and W M Ross ldquoRelationshipof stalk morphology and chemical composition to lodgingresistance in sorghumrdquo Crop Science vol 17 no 4 p 609 1977
[40] P K Das B C Jena and N Nayak ldquoCharacter association andpath analysis of sugar yield in sugarcanerdquo Indian Sugar vol 46no 10 pp 805ndash808 1997
[41] K J Sreekumar M Kuriakose M Thomas D Alexander andS Santhakumari ldquoVariability heritability and correlation onstudies on the yield and quality characters of sugarcanerdquo IndianSugar vol 44 no 4 pp 243ndash249 1994
[42] J R Harlan J M de Wet and E G Price ldquoComparativeevolution of cerealsrdquo Evolution vol 27 no 2 pp 311ndash325 1973
[43] A Elings ldquoDurum wheat landraces from syria II patterns ofvariationrdquo Euphytica vol 54 no 3 pp 231ndash243 1991
[44] K Tato ldquoRainfall in Ethiopiardquo Ethiopian Geographical Journalvol 2 pp 28ndash36 1964
[45] A B L Stemler J R Harlan and J M J deWet ldquoThe sorghumsof ethiopiardquo Economic Botany vol 31 no 4 pp 446ndash460 1977
[46] P Annicchiarico L Pecetti and A B Damania ldquoRelationshipsbetween phenotypic variation and climatic factors at collectingsites in durum wheat landracesrdquo Hereditas vol 122 no 2 pp163ndash167 1995
[47] A B Damania L Pecetti C O Qualset and B O HumeidldquoDiversity and geographic distribution of adaptive traits inTriticum turgidum L (durum group) wheat landraces fromTurkeyrdquoGenetic Resources and Crop Evolution vol 43 no 5 pp409ndash422 1996
[48] A Ayana and E Bekele ldquoGeographical patterns of morpho-logical variation in sorghum (Sorghum bicolor (L) Moench)germplasm from Ethiopia and Eritrea quantitative charactersrdquoEuphytica vol 115 no 2 pp 91ndash104 2000
[49] D S Falconer Introduction to Quantitative Genetics JohnWilley and Sons Inc New York NY USA 3rd edition 1993
[50] R S Thorpe ldquoBiochemical analysis of geographical variationand racial affinitiesrdquo Biology Review vol 51 pp 407ndash452 1976
[51] M Farooq ldquoMorphological and agricultural characteristicsof sugarcane in relation to mechanized agriculture at NARCIslamabadrdquo Portu Rico Journal vol 1 pp 51ndash61 1989
[52] M Akhtar N N Elahi and M Ashraf ldquoEvaluation of exoticsugarcane varieties for agronomic characters and productivityrdquoPakistan Journal of Biological Sciences vol 4 no 1 pp 37ndash402001
[53] F R Piscitelli ldquoPrincipals caracteres exomorfologicos de cul-tures de cana de azucarrdquo TUC Iindustraly Agricos vol 7 pp 49ndash57 1994 (Spanish)
[54] A H D Brown and B S Weir ldquoMeasuring genetic variability inplant populationsrdquo in Isozyme in Plant Genetics Breeding S DTanksley and T J Orton Eds part A Elsevier Science 1983
[55] R Balakrishnan N V Nair and T V Sreenivasan ldquoA methodfor establishing a core collection of Saccharum officinarum Lgermplasm based on quantitative-morphological datardquo GeneticResources and Crop Evolution vol 47 no 1 pp 1ndash9 2000
[56] C A Nestor and B Q Reynaldo Sugarcane Variety Improve-ment in Southeast Asia and the Pacific for Enhanced andSustainable Productivity ndash Germplasm and Disease IndexingComponents Institute of Plant Breeding (IPB) College ofAgriculture (CA) UP Los Banos (UPLB) 2003
[57] A Maqbool J Muhammad and A Sagheer ldquoAgronomic traitsand morphological characteristics of some exotic varieties ofsugarcanerdquoPakistan Journal of Agricultural Research vol 19 no4 2006
Most of the sampled germplasm had acceptable levels of juicerefractometer readings (expressed in degree brix) relative tothe history of the ages of the samples (Supplemental Table 1)The number of harvestablesaleable cane stalks obtained inplant cane and ratoon crops and the number of harvestableratoon crops showed variation region to region and zone tozone and sometimes district (ldquoweredardquo)-wise There is alsovariation in maturity time of the clones both in plant caneand in ratoon crops (Supplemental Table 2)These are mainlydepending on the variety soil condition cultural practicesand climatic conditions
The genotypes collected from the respective zones inthe study regions were recorded by their local namesSometimes similar varieties bear different names at differentlocalities and vice versa Farmers mentioned a broad rangeof sugarcane landraces that had been grown in the areasand maintained for generations Some of the landraces werecommonly recognized by most farmers within and acrosszones of the respective regions whereas some were rarevarieties known only by few farmers The large number oflandraces observed during the current study demonstratesthe existence of diverse genetic resources of sugarcane inEthiopia The diversity could have evolved presumably due todiverse climatic conditions low input management systemshigh pests and disease pressure and continuous selectionby farmers Smallholder farmers face diverse environmentstresses and have multiple production objectives that affectselection of genotypes [24] There were no formally releasedimproved sugarcane varieties grown by farmers in the studyregions except in few places This was mainly observedin SNNPR at Wolayta and Kembata-Tembaro zones Thesevarieties have been informally introduced to these areas byseasonal workers employed at the sugar estates especially atWonji and Metehara
The collected landraces would serve to broaden thegenetic base of the available sugarcane germplasm Thesugar industry in Ethiopia is currently establishing sugarcanebreeding programTherefore collection and efficient charac-terization of germplasm are vital to strengthen the breedingprogram
311 Physical and Chemical Properties of Soils of Sugar-cane Germplasm Collection Areas Table 2 summarizes thephysical and chemical properties of soils in the sugarcane
production fields of surveyed areas Sugarcane productionby smallholder farmers in the surveyed areas is conductedacross the seasons where harvesting and planting are done aslong as there is available moisture Therefore the soil samplesrepresent the sugarcane production season and the altitudinalranges of the respective areas
Generally the organic carbon content of soils in sugarcaneproducing areas across the country is low to very low (2-4lt2) Amhara Gambella SNNP Somali and Tigray Regionsare in a very low category (lt2) Similarly across regions thetotal N content is low except Harari and Oromia regionswhere the average total N is medium This problem shouldbe addressed either with augmenting the soil with inorganicfertilizers or through appropriate management practices likecrop residue management Available phosphorus is high inAmhara Gambella and SNNP Regions whereas moderatevalues were observed in Harari Oromia and Tigray RegionsGenerally the level of phosphorus in soils of sugarcaneproducing areas in these regions is adequate In Benshangul-Gumz and Somali Regions low level of phosphorus wasrecorded which should be addressed with due attentionAvailable K is sufficient for normal sugarcane growth in soilsof all regions Relatively lower values of K were recorded insoils of Harari and Tigray Regions It should be noted thatorganic carbon EC PH and variability of P andK are affectedby rainfall and temperature that are seasonal and the valuesrecorded refer to the seasonwhen the soil samples were takenGenerally Ethiopian soils are deficient in various essentialnutrients like boron nitrogen phosphorus potassium sulfurzinc and copper although severity differs from region toregion [25] Particularly the loss of P and N resulting fromthe use of dung and crop residues for fuel is demonstrated tobe equivalent to the total amount of commercial fertilizer use
Across regions in Ethiopia generally clay type of soildominates followed by silt except in Gambella Region wherethe dominant soil texture was sand Soil management optionsrelated to clay soil may address soil related problems insmallholder sugarcane growing areas across Ethiopia
32 Diversity of Sugarcane Landraces
321 Quantitative Characters
Univariate Statistics Analysis of variance depicted highlysignificant differences (p lt 001) between accessions pooled
6 Advances in Agriculture
over the regions for the 21 characters of the 211 sugarcaneaccessions studied and significant difference for the 6 regionsof origin for 6 characters (Table 3) The results suggestedthe occurrence of significant phenotypic variation betweenthe accessions as a whole Significant variations of differentsugarcane stalk and juice quality characters was also reportedin similar studies conducted elsewhere [26 27] Region-wise partitioning of the variance indicated significant within-region differences (p lt 005) among the populations withinalmost all regions for the characters NOI IL SH SD LLLW LA Brix Pol Purity SR MSCHA SCW CYHAand SY for 8 characters within Tigray for TC4MAP andTC5MAP within Oromiya and SNNPR Brix10MAP showedno significant difference within all regions
In general within-region variation was greater for stalkdiameter single cane weight millable stalk count and caneand sugar yield than for other characters for all the regionsAssuming that a significant portion of the phenotypic vari-ation is genetic it would be possible to make selection forany of this group of characters within a particular regionIt was apparent that the variance between accessions pooledover regions was greater than between regions Therefore inorder to sample the variation effectively it would be necessaryto sample the variable populations from different localitiesin a region The study in [28] on genetic diversity amongmain land USA sugarcane cultivars showed high geneticdiversitywithin populations of different regions than betweenpopulations
Duncanrsquos multiple range testing for regional means for allthe characters is shown in Table 4 Relatively much differenti-ationwas observed for stemdiameter and juice purity percentcompared to other parameters Important yield componentslike MSCHA SCW SH and SD have shown higher valuesin Tigray SNNPR and Oromiya regions High cane andsugar yield were recorded for accessions in Tigray regionfollowed by SNNPR Oromiya and introduction which werestatistically at par Statistically similar lower values of caneand sugar yield were recorded in Amhara and Benshangulcollections
Higher value of sucrose percent in cane was also observedfor accessions from Tigray and the lowest value was inAmhara accessions while other regions were statisticallysimilar Generally for most of the characters high valuewas obtained from accessions in Tigray region followed bySNNPROromiya and introduction Therefore the materialsfrom these regions can be used for selection of cane and sugaryield per se and in breeding program The local accessions inmany of the regions showed superiority over the introducedcommercial varieties currently in production for most of thecharacters including cane and sugar yield (Table 4)This maybe due to the adaptation and coevolution of the materialswith different stresses for long period of time in the countryrsquosagroecology These locally adapted genes conferring yieldadvantage could be harnessed through strategic crossing andselection for improvement of the crop Superiority of locallandraces over introduced varieties was reported [29 30]
The range of variation of the accession means establishedwide variation between the regions and the accessions withinthe regions for the characters studied (Table 5) Accordingly
the maximum score was 41 times the minimum for sproutcount one month after planting 32 for sprout count twomonths after planting 25 for tiller count four months afterplanting 22 for sugar yield 20 for cane yield 12 for millablestalk count 8 for stalk count 10 months after planting 7for single cane weight 6 for tiller count five months afterplanting and 2 for other yield components and sugar qual-ity parameters except hand refractometer brix reading tenmonths after planting and juice purity percent which scored1 The same trend was observed between accessions withina particular region However accessions within AmharaBenshangul-Gumz and introduction showed wider ranges ofvariation than those accessions within other regions for themajority of the characters The wide range of geographic andclimatic features of the regions in Ethiopia and the originaldifferentiation and further farmers selection for productionniches and uses for hundreds of years may have resultedin the accessions to possess a tremendously high degree ofmorphological variation The reason for introduced varietiesshowing high variation also could be due to broader geo-graphic spectrum fromwhere theywere initially acquired anddifference in their parental source and genealogical historyIt was demonstrated even with these limited numbers ofsamples (15 genotypes) that introduction of genetic materialshas practically broadened the genetic base of the sugarcanegermplasm for the breeding program
Coefficient of variation measured as the ratio of standarddeviation to the corresponding overall mean and expressedas percentage is useful to compare different charactersmeasured in different units It is also useful to compare samecharacter in different groups of populations with differentsample size mean and variance or different characters indifferent populations In the current study high coefficientsof variation were observed between regions and within eachregion for sprout as well as tiller counts in the monthsthe data was recorded stalk count 10 months after plant-ing and millable stalk count at harvest cane and sugaryield single cane weight and leaf area (Table 6) Thoughaccessions from a particular region were more variable fora specific character compared to other regions accessionsfromAmhara Benshangul-Gumz introduction and SNNPRwere more variable than accessions from other regions Thisresult demonstrated the tremendous variability of sugar-cane germplasm from these regions The accessions fromTigray and Oromiya had low coefficients of variation formany characters indicating relatively high within-regionuniformity The high coefficients of variation observed formost of the characters agreed well with those reportedin [31] for indigenous sugarcane of Brazil Similar resultswere also reported in sorghum [32] The different levels ofregional variability of a particular character could be due todifferences in forces of selection andor differences in theintensity of a particular selecting force It could also be dueto random effect and reduced sample parents of parentalmaterials moved differently by various human populationmigration and the interaction of these with Darwinianforces
Results of this study suggested the presence of wide rangeof variations of characters determined by univariate statistics
This is in agreement with previous studies on sugarcanegermplasm [33ndash35]
This is the first study to report information on geo-graphical pattern of variation in Ethiopian local sugarcanegermplasm which has been lacking so far Our results pro-vided experimental evidence on occurrence of geographicalvariation and significant within-region variation where it washigh in the regions such as Amhara Benshangul-Gumz andSNNPR
The overall patterns of similarity or difference betweenregions seemed to depend on environmental factors such asrainfall temperature length of growing season and altitudeWide variability of millable stalk count at harvest singlecane weight and plant height was observed among regionswhere these characters demonstrated high correlation withcane and sugar yield (Table 7) Therefore breeders can utilizeaccessions of regions showing variability for these charactersin selection programs and to design breeding strategies toproduce varieties with best commercial merits
Bivariate Statistics Correlation coefficients worked out onthe 21 quantitative characters are shown for the entire data(Table 7) between regions (Table 8) and within regions(data not shown) Results of the association studies for theentire data showed that cane and sugar yield had highlysignificant positive correlation with most of the quantitativestalk characters and juice quality parameters However theassociation of tiller counts 4 and 5 months after plantingstalk count 10 months after planting millable stalk countat harvest single cane weight and plant height with caneand sugar yield was stronger suggesting these componentcharacters asmain contributing factor to cane and sugar yieldIn fact sugar yield is the product of cane yield and sucrose per-cent cane surely being influenced with components affectingcane yield Similar trend was reported by earlier sugarcaneworkers [36 37] In sugarcane the cane and sugar yields areconsidered to be the complex characters The information onthe phenotypic interrelationship of cane yield and sugar yieldwith their component character per se would be of immensehelp to sugarcane breeders
Stem diameter had significant positive association withboth cane and sugar yield indicating the significance ofthis trait in improving both cane and sugar yields Similarreports have beenmade by earlier sugarcane workers [37 38]Stalk diameter was also positively and significantly correlatedwith plant height It is one of the traits that are related tolodging resistance [39] and the positive associations withheight would help in reducing the chance of lodging as heightincreases Among the physiological attributes included in thestudy leaf area had highly significant positive associationwith cane yield Similar observation was also made in [37]Sucrose percent which is an important juice quality attributehad highly significant positive association with sugar yieldand simultaneously cane yield also had highly significantassociation with sugar yield Based on the magnitude ofcorrelation coefficient values it can be said that cane yield ismuch more important than sucrose percent in determiningthe sugar yield and the same has also been reported by othersugarcane workers [37 40] Similar to observations made in
[37 41] results of the current study showed that all the qualityparameters like sucrose percent pol percent brix percentand purity percent were significantly correlated in positivedirection This indicated any of these juice quality traitscould be considered for selection leading to the simultaneousimprovement in the remaining quality traits
Ten of the 21 characters also showed significant positivecorrelations with altitude of the collection sites (Table 7) Asdescribed in [42] ecological characteristics have influencedthe genotypic constitution of landraces during domesticationand hence a relationship exists between the agroecology inthe collection site and the morphological characteristics ofthe landraces Thus positive correlation between collectionsite variables and plant characteristics would imply that thevariation between accessionsmay be related to agroecologicalvariations among the collection sites [43]
The correlation coefficient between number of internodesand altitude was negative and significant indicating thatother environmental factors (other than altitude) andornonenvironmental factorsmight account for the variation forthis particular character Since temperature decreases withan increase of altitude in Ethiopia [44 45] it is more likelythat temperature has exerted strong selection pressure innumber of internodes in the genotypes Information on therelationships between environmental factors of the collect-ing sites and morphophysiological variation of germplasmcould enhance the understanding of evolutionary adaptivepatterns which could assist germplasm collectors and users[46]
Character associations between region and within regionalso followed similar fashion like that of the entire dataNumber of sprout tiller and stalk counts and sugar qualityparameters showed significant positive correlations with caneand sugar yield
Though not significant and strong the correlation of leaflength and width and leaf area (leaf area = leaf length Xleaf width X 075) with cane and sugar yield was negativein the interregion correlations This kind of relationship wasalso exhibited between these leaf characteristics and yieldcomponents like tiller and stalks counts plant height andsingle cane weight and stalk diameter and sugar qualityparameters This may have been attributed to the variationof temperature with altitude and other environmental factorsacross the regions resulting in such type of relationshipSuch relationship also calls for the need to have enough dataon environmental variables of the collection sites such astemperature and rainfall if germplasm collection is requiredtomeet its objectives effectively and efficiently Unfortunatelydata on such useful environmental variables like temperatureand rainfall are lacking in the passport data of the currentEthiopian sugarcane germplasm collections The significanceof environmental factors of the collection sites as veryimportant determinants in structuring morphophysiolog-ical variations was reported in tetraploid wheat [22 4748]
Association of characters among yield its componentsand other economic traits is important for the interpretationof the patterns of variation and making selection in breed-ing program and combining several desirable attributes It
suggests the advantage of a scheme of selection for morethan one character at a time The correlation between char-acters may arise from linkage or from developmental geneticinteractions with orwithout a purely phenotypic componentIt could arise also due to differential phenotypic plasticityof characters themselves Within the limit of experimentalerror and environmental effects high correlation coefficientsbetween characters may show that the characters sharesome common element of genetic control (ie pleiotropylinkage) between genes or else from independently controlledcharacters responding similarly to geographic variation inselection pressures [49] The interregion correlation coef-ficient between the characters measures the consistency oftheir patterns of regional variation while the intraregioncorrelation coefficient measures the association arising fromgenetic factors but is not affected by regional variation[50] Since this study showed significant positive correlationsintraregionally for some character combinations it wouldseem that common genetic control might be playing a role inbringing about correlations between the various charactersIt appeared that different response to regional variationwas playing a greater role than different genetic controlas shown by the many more significant and moderate tohigh correlation coefficients intraregionally than interregion-ally
322 Qualitative Characters The frequency distribution forthe 16 qualitative characters of germplasm samples by regionsin Ethiopia and of the exotic accessions is shown in Supple-mental Table 5 For the purpose of avoiding redundancy andimproving readability the discrete characters are presented bygrouping related traits together
BudCushion andRelativeDegree of BudExtension Evaluationof the presence and absence of bud cushion revealed thatgreat proportion (70) of sugarcane landraces from Ethiopiado not have bud cushion Majority of the sample germplasmin all regions belong to this phenotypic class Approximatelyone-half (53) of the collection have their bud extendedabove the growth ring followed by those (41) having budtouching the growth ring
Relative Bud Shape Among the phenotypic classes for thischaracter tall deltoid was dominant (31) followed by ovate(17) and narrow ovate (12) While local accessions inmost regions had tall deltoid bud shape 50 of the intro-duced accessions belong to two phenotypic classes roundwith central germpore (30) and 20 having pentagonalbud shape Equal proportions (19) of the samples fromBenshangul-Gumz were Ovate and Ovate with emarginatebasal wing
Canopy Structure Relative Plant Erectness Colour of theLeaves and Colour of the Exposed Rind Out of the eightphenotypic classes observed compact tip droopy was themost frequent (35) Open semidroopy and open droopyclasses had fairly equal distribution With regard to relativeplant erectness seventy-five percent of the germplasm werefound to fall in twophenotypic classes erect (41) and almost
erect (34) for the whole region and country Most of theaccessions (90) showed light green or green leaf colourbut 20 from introduced collections were greenish yellowAmong phenotypic classes for colour of the exposed rindbrownish yellow yellow brown and yellowish green were themost frequent with fairly equal distribution
Relative Shape of Dewlap Double crescent had larger fre-quency (22) while squarish descending and flaring typeof dewlap shape had fairly equal proportions Seventeenpercent of germplasm from Tigray region had squarishdeltoid dewlap shape
Relative Degree of Internode Alignment and Internode ShapeForty-four percent of the sample germplasm were slightlyzigzag while there was almost equal frequency for straightand zigzag internode alignment for the entire region andcountry Straight or nearly straight (slightly zigzag) internodealignment of cane stalks is a very important character formechanized farming and postharvest handling in sugarcane[51] Forty percent of the collections were observed havingconcave-convex type of internode shape while 29 wascylindrical and 14 conoidal Other classes had fairly equalproportion
Type of Auricle and Relative Shape of Ligule An equalproportion of the germplasm totally constituting 84 ofthe collection belong to four phenotypic classes namelythose with no auricle and others with transitional and shortand lanceolate auricles Sixteen percent of the collectionsalso showed equal frequency for the remaining classesBenshangul-Gumz and Tigray regions had each equal 13accessions with falcate type of auricle Sixty-seven percentof the collections belong to three ligule shape classes cres-cent with lozenge (27) broad-crescent (21) and linear-crescent (19) The germplasm had almost equal proportionfor the other phenotypic classes of this character Nineteenpercent of the accessions fromBenshangul-Gumz region haddeltoid shape of ligule
Stalk Corky Cracks Corky Patches Growth Cracks and BudGroove Eighty-nine percent of the germplasm across regionshad no stalk corky cracks with only eleven percent exhibitingthis character All the germplasm in Gambella Harari andSomali had no stalk corky cracks On the other hand eighty-one percent of the germplasm for the whole region showedthe presence of stalk corky patches All the local germplasmin Benshangul-Gumz Gambella and Tigray had stalk corkypatches whereas 75 of the germplasm from Harari Region67 from Somali Region and 24 from Oromia Regionhad no stalk corky patches In the entire region 75 ofthe germplasm had no stalk growth cracks with only 25showing this character Sixty-three percent of the germplasmwere found to have bud groove Howevermost of the samplesin Harari (75) Amhara (63) and Oromia (53) Regionshad no bud groove
The results of this study indicated the wide distributionof phenotypic classes for the characters considered whichindicates the existence of different races and combination
Advances in Agriculture 15
of races in local genotypes of different regions in EthiopiaThis diversity of germplasmbased onphenotypicmarkers canhelp sugarcane breeders in identifying the identity of geno-types as well as maintaining genetic diversity Differencesin morphological characters of different sugarcane varietieshave been reported [52] Piscitelli [53] showed importantexomorphological qualitative characters of sugarcane varietythat are not influenced by environmental factors and thus canbe used as a selection tool in any breeding program
The amount of phenotypic diversity estimates based onthe ShannonndashWeaver diversity index (H1015840) are shown inTable 9 The 16 characters differed in their distribution aswell as the amount of variation Individual traits showed adifferent pattern of variation among accessions Estimatesof H1015840 for individual characters varied from 049 for stalkcorky cracks to 100 for bud groove with an overall meanof 080 Most of the characters were highly polymorphicwhereas relative plant erectness and presence or absenceof stalk corky patches scored moderate values The highdiversity values for the characters showed a wide variabilityamong genotypes According to [54] this index is also usedin genetic resource studies as a convenient measure of bothallelic richness and allelic evenness when using geneticdata
The highest diversity computed was on presence orabsence of bud groove with a diversity index of 100indicating that this character is very variable in the localsugarcane germplasm On the other hand the descriptorcomputed with the lowest diversity index value (049) wasthe presence or absence of stalk corky cracks indicatingthat this character is selected in the development of thegenotypes In agreement with the present study [55] workingon S officinarum accessions from the world collection ofsugarcane germplasm obtained a low index of 025 and 035for presence or absence of stalk corky patches and cracksrespectively
Their report also showed high diversity index values forinternode alignment growth cracks internode shape budcushion bud extension bud groove and ligule shape Sim-ilarly [56] experimenting on field collections of sugarcaneaccessions in Philippine reported high ShannonndashWeaverdiversity for stalk growth cracks bud cushion stalk corkypatch stalk corky cracks bud extension and auricle outershape Unlike the present study they have reported high indexfor stalk corky cracksMedium index values were reported forstalk internode shape and low index values for ligule shapeand bud shape High variability of sugarcane varieties fordewlap shape was also reported in [57]
The H1015840 pooled across characters by region ranged from079 in Oromiya to 085 in Tigray with an overall aver-age of 081 (Table 8) Most of the regions showed highShannonndashWeaver diversity index The populations that hadthe highest H1015840 were that of Tigray and introduction The H1015840averaged over regions for different characters was found torange from 044 for presence or absence of stalk corky cracksto 094 for relative degree of internode alignment and budgroove with an overall average of 081 (Table 9)
Assessment of the ShannonndashWeaver index also showedsensitivity to both the number of descriptor classes and the
distribution within phenotypic classes in each region Amongregions the mean values for the ShannonndashWeaver index arenot significantly different but differences were found whenindividual characters were considered These observationssuggest that some regions have high diversity for particulartraits while in other regions selective pressures might havereduced the variation to certain genotypes
4 Conclusions
The large number of landraces observed during the cur-rent study demonstrates the existence of diverse geneticresources of sugarcane in Ethiopia The landraces wouldserve to broaden the genetic base of the available sugarcanegermplasm Generally distribution of the overall phenotypicdiversity among sugarcane landraces in Ethiopia is unevenOur results showed that there was a wide range of variationthat existed in the local sugarcane landraces studied both atregional and within region levels Selection and differenti-ation of types might have taken place along a geographicalpattern perhaps associated with climate and use The presentstudy contributes to updating sugarcane descriptors adoptedfrom USDA-ARS as well as Bioversity passport data forthe future collection and evaluation As Ethiopian SugarCorporation is in the course of launching breeding programthe observed substantial variation of landraces would enablesugarcane breeders to design and practice breeding andselection programs to improve the crop
Data Availability
The data used to support the findings of this study areavailable from the corresponding author upon request
Conflicts of Interest
The authors declare that they have no conflicts of interest
Acknowledgments
The authors are grateful for the financial grant of SugarCorporation of Ethiopia Many thanks are due to EthiopianBiodiversity Institute (EBI) for its technical advice duringcollection of the local sugarcane genotypes Sincere thanksare forwarded to Ethiopian smallholder farmers who pre-served sugarcane germplasm and shared their knowledgeand experience Regional Zonal and District (ldquoWeredardquo)Agriculture Bureaus across Ethiopia are acknowledged fortheir support during the study
Supplementary Materials
These supplementary materials contain different tables sup-porting the results of the study These include passport dataof sugarcane germplasm collected and related informationon the materials local sugarcane germplasm collected andused formorphological diversity study introduced sugarcane
16 Advances in Agriculture
Table9Sh
anno
n-WeaverD
iversityIndices(H1015840)a
ndmeanandsta
ndarderror(plusmnS
E)of
land
racesineach
region
basedon
16discretecharacters
Characters
Amha
raBe
nsha
ngul-G
umz
Introd
uced
Oromia
SNNP
Tigray
ELb
Total
Presence
orabsenceo
fbud
cushion(BUDCUSH
ION)
083
095
095
089
082
098
090
088
Relativ
edegreeo
fbud
extension(BUDEX
TEND)
077
079
093
074
072
087
080
078
Relativ
ebud
Shape(BU
DSH
APE
)078
096
089
081
085
078
084
078
Canop
ystructure(C
ANOPY
)089
098
095
081
078
096
089
084
Relativ
eshape
ofdewlap(D
EWLA
PSHAP)
087
094
092
090
082
096
090
086
Relativ
eplant
erectness(ER
ECT)
080
082
050
067
071
088
073
064
Relativ
edegreeo
finterno
dealignm
ent(IN
ALIGN)
096
076
098
097
098
100
094
098
Relativ
einterno
deshape(IN
SHAPE
)086
069
095
077
083
091
083
082
Color
oftheleaves(LE
AFC
OLO
R)087
071
090
064
074
071
076
076
Type
ofou
tera
uricle(A
URI
CLE
OUT)
080
092
087
079
083
093
086
082
Relativ
eshape
ofligule(LIGSH
APE
)086
094
082
085
090
083
087
083
Presence
orabsenceo
fstalkcorkyc
racks(ST
ALK
CORK
C)
044
054
035
055
051
025
044
049
Presence
orabsenceo
fstalkcorkyp
atches(STA
LKCORK
P)066
000
072
079
059
087
060
069
Presence
orabsenceo
fstalkgrow
thcracks
(STA
LKCRA
CK)
066
100
088
062
088
099
084
080
Color
ofthee
xposed
rind(R
INDCOLE
)085
096
089
084
086
076
086
085
Presence
orabsenceo
fbud
groo
ve(BUDGRO
OVE)
095
100
072
100
100
099
094
100
Means
080plusmn0
03
081plusmn0
06
083plusmn0
04
079plusmn0
03
080plusmn0
03
085plusmn0
05
081plusmn0
03
080
b EL=
averageo
fentire
locatio
n
Advances in Agriculture 17
varieties used for morphological diversity study and fre-quency distribution for 16 qualitative characters in sugarcaneby regions in Ethiopia and country of sources of introducedgenotypes (Supplementary Materials)
References
[1] ldquoSpreading the word about clean solutions from sugarcanrdquoAccessed 04072018 httpwwwsugarcaneorg
[2] Central Statistics Agency (CSA) ldquoAgricultural sample surveyArea and production of major crops private peasant holdingsrdquoStatistical Bulletin Addis Ababa Ethiopia 2017
[3] T Esayas Exploration and Collection Characterization GeneticDiversity Analysis and Association of Traits for Yield and YieldComponents of Sugarcane (Saccharumspp) in Ethiopia [MasterThesis] Haramaya University Dire Dawa Ethiopia 2014
[4] J R Harlan Crops and Man American Society of AgronomyCrop Science Society of America Madison WI USA 2ndedition 1992
[5] R W Allard Principles of Plant Breeding JohnWilley and SonsNew York NY USA 2nd edition 1999
[6] B T Roach and J Daniels ldquoA review of the origin and improve-ment of sugarcanerdquo in Proceedings of Coper sugar InternationalSugarcane Workshop pp 1ndash32 1987
[7] P P Patil S S Patil and D U Patel ldquoGenetic diversity in sug-arcane (Saccharum spp Complex)rdquo Journal of Pharmacognosyand Phytochemistry vol 6 no 6 pp 570ndash573 2017
[8] R B Singh B Singh andR K Singh ldquoStudy of genetic diversityof sugarcane (Saccharum) species and commercial varietiesthrough TRAP molecular markersrdquo Indian Journal of PlantPhysiology vol 22 no 3 pp 332ndash338 2017
[9] R W Allard ldquoPopulation structure and sampling methodsrdquo inGenetic Resources in PlantsmdashTheir Exploration and Conserva-tion O H Frankel and E Bennett Eds IBP Handbook no 11pp 97ndash107 FA Davis Company Philadelphia PA USA 1970
[10] D R Marshall and A H D Brown ldquoOptimum samplingstrategies in genetic conservationrdquo inCrop Genetic Resources forToday and Tomorrow O H Frankel and J G Hawkes Eds pp53ndash80 Cambridge University Press 1975
[11] J G Hawkes Crop Genetic Resources Field Collection ManualPudoc Wagnningen Netherland 1980
[12] J G Hawkes The Diversity of Crop Plants Harvard UniversityPress London England 1983
[13] B P K Yerima Manual for Good Laboratory Practice Foodand Agriculture Organization of United Nation Addis AbabaEthiopia 1993
[14] A Walkley and I A Black ldquoAn examination of the degtjareffmethod for determining soil organic matter and a proposedmodification of the chromic acid titrationmethodrdquo Soil Sciencevol 37 no 1 pp 29ndash38 1934
[15] S R Olsen and L E Sommers ldquoPhosphorusrdquo in AgronomyMonograph A L Page et al Ed Methods of soil analysis part2 pp 403ndash430 ASA and SSSAMadisonWI USA 2nd edition1982
[16] R H Bray and L T Kurtz ldquoDetermination of total organic andavailable forms of phosphorus in soilsrdquo Soil Science vol 59 no1 pp 39ndash46 1945
[17] S Sahlemedhin andB Taye ldquoProcedures for soil and plant anal-ysisrdquo Tech Rep 74 National Soil Research Center EthiopianAgricultural Research Organization 2000
[18] F C Stickler S Weaden and AW Pauli ldquoLeaf area determina-tion in grain sorghumrdquo Journal of Agronomy and Crop Sciencevol 53 pp 187-188 1961
[19] Munsell Color ChartThe Munsell Book of Color Matte Collec-tion Munsell Color Chart NewWindsor NY USA 1990
[20] H O Hartley ldquoThe maximum F-ratio as a short cut test forheterogeneity of variancesrdquo Biometrika vol 37 pp 308ndash3121950
[21] L Pecetti P Annicchiarico and A B Damania ldquoBiodiversity ina germplasm collection of durumwheatrdquo Euphytica vol 60 no3 pp 229ndash238 1992
[22] L Pecetti and A B Damania ldquoGeographic variation intetraploid wheat (Triticum turgidum ssp turgidum convardurum) landraces from two provinces in Ethiopiardquo GeneticResources and Crop Evolution vol 43 no 5 pp 395ndash407 1996
[23] S K Jain C O Qualset G M Bhatt and K K Wu ldquoGeo-graphical patterns of phenotypic diversity in a world collectionof durumwheatsrdquoCrop Science vol 15 no 5 pp 700ndash704 1975
[24] K K Nkongolo L Chinthu M Malusi and Z Vokhiwa ldquoPar-ticipatory variety selection and characterization of sorghum(Sorghum bicolour (L) Moench) elite accessions from Malaw-ian gene pool using farmer and breeder knowledgerdquo AfricanJournal of Agricultural Research vol 3 pp 273ndash283 2008
[25] M B Semira ldquoInnovative methods and metrics for agricul-ture and nutrition actions soil quality and human nutritionin Ethiopiardquo Leverhulme Center for Integrative Research onAgriculture and Health (LCIRAH) 2017
[26] I A Khan A Khatri M A Siddiqui G S Nizamani and SRaza ldquoPerformance of promising sugarcane clone for yield andquality traits in different ecological zones of Sindhirdquo PakistanJournal of Botany vol 36 no 1 pp 83ndash92 2004
[27] M F Perera M E Arias D Costilla et al ldquoGenetic diversityassessment and genotype identification in sugarcane based onDNAmarkers andmorphological traitsrdquo Euphytica vol 185 no3 pp 491ndash510 2012
[28] C G Neil M Kay and C C Jack ldquoDiversity among mainlandUSA sugarcane cultivars examined by SSR genotypingrdquo Journalof the American Society of Sugar Cane Technologists vol 29 pp36ndash52 2009
[29] S SchenckMWCrepeauK KWu P HMooreQ Yu andRMing ldquoGenetic diversity and relationships in native HawaiianSaccharum officinarum sugarcanerdquo Journal of Heredity vol 95no 4 pp 327ndash331 2004
[30] R Gebretsadik H Shimelis M D Laing P Tongoona andN Mandefro ldquoA diagnostic appraisal of the sorghum farmingsystem and breeding priorities in Striga infested agro-ecologiesof Ethiopiardquo Agricultural Systems vol 123 pp 54ndash61 2014
[31] L E V Cesar A T Bruzi J A R Nunes et al ldquoGeneticdivergence of sugarcane varieties collected in the region ofLavras Minas Gerais Brazilrdquo Genetics and Molecular Researchvol 14 no 4 pp 14043ndash14049 2015
[32] C Grenier P J Bramel J A Dahlberg et al ldquoSorghums of theSudan analysis of regional diversity and distributionrdquo GeneticResources and Crop Evolution vol 51 no 5 pp 489ndash500 2004
[33] N V Nair R Balakrishnan and T V Sreenivasan ldquoVariabilityfor quantitative traits in exotic hybrid germplasm of sugarcanerdquoGenetic Resources and Crop Evolution vol 45 no 5 pp 459ndash464 1998
[34] R Nagarajan S Alarmelu and R M Shanthi ldquoStudies onvariation in interspecific hybrids of Saccharumrdquo Sugar Tech vol2 no 3 pp 42ndash46 2000
18 Advances in Agriculture
[35] D N Kamat and J R P Singh ldquoVariability in Sugarcane underrain fed conditionrdquo Sugar Technology Short Communicationsvol 3 no 1-2 pp 65ndash67 2001
[36] B S Rishipal A S M Chaudhary and S P Kadian ldquoStudieson character interrelationship in clonal generation of sugarcane(Saccharum complex hybrids)rdquo Indian Sugar vol 47 no 11 pp907ndash911 1998
[37] S Thippeswamy S T Kajjidoni P M Salimath and J V GoudldquoCorrelation and path analysis for cane yield juice quality andtheir component traits in sugarcanerdquo Short Communication vol5 no 1-2 pp 65ndash72 2003
[38] P S Verma S Pal and N K Karma ldquoGenetic variability andcorrelation studies in sugarcanerdquo Indian Sugar vol 49 pp 125ndash128 1999
[39] H A Esechie J W Maranville and W M Ross ldquoRelationshipof stalk morphology and chemical composition to lodgingresistance in sorghumrdquo Crop Science vol 17 no 4 p 609 1977
[40] P K Das B C Jena and N Nayak ldquoCharacter association andpath analysis of sugar yield in sugarcanerdquo Indian Sugar vol 46no 10 pp 805ndash808 1997
[41] K J Sreekumar M Kuriakose M Thomas D Alexander andS Santhakumari ldquoVariability heritability and correlation onstudies on the yield and quality characters of sugarcanerdquo IndianSugar vol 44 no 4 pp 243ndash249 1994
[42] J R Harlan J M de Wet and E G Price ldquoComparativeevolution of cerealsrdquo Evolution vol 27 no 2 pp 311ndash325 1973
[43] A Elings ldquoDurum wheat landraces from syria II patterns ofvariationrdquo Euphytica vol 54 no 3 pp 231ndash243 1991
[44] K Tato ldquoRainfall in Ethiopiardquo Ethiopian Geographical Journalvol 2 pp 28ndash36 1964
[45] A B L Stemler J R Harlan and J M J deWet ldquoThe sorghumsof ethiopiardquo Economic Botany vol 31 no 4 pp 446ndash460 1977
[46] P Annicchiarico L Pecetti and A B Damania ldquoRelationshipsbetween phenotypic variation and climatic factors at collectingsites in durum wheat landracesrdquo Hereditas vol 122 no 2 pp163ndash167 1995
[47] A B Damania L Pecetti C O Qualset and B O HumeidldquoDiversity and geographic distribution of adaptive traits inTriticum turgidum L (durum group) wheat landraces fromTurkeyrdquoGenetic Resources and Crop Evolution vol 43 no 5 pp409ndash422 1996
[48] A Ayana and E Bekele ldquoGeographical patterns of morpho-logical variation in sorghum (Sorghum bicolor (L) Moench)germplasm from Ethiopia and Eritrea quantitative charactersrdquoEuphytica vol 115 no 2 pp 91ndash104 2000
[49] D S Falconer Introduction to Quantitative Genetics JohnWilley and Sons Inc New York NY USA 3rd edition 1993
[50] R S Thorpe ldquoBiochemical analysis of geographical variationand racial affinitiesrdquo Biology Review vol 51 pp 407ndash452 1976
[51] M Farooq ldquoMorphological and agricultural characteristicsof sugarcane in relation to mechanized agriculture at NARCIslamabadrdquo Portu Rico Journal vol 1 pp 51ndash61 1989
[52] M Akhtar N N Elahi and M Ashraf ldquoEvaluation of exoticsugarcane varieties for agronomic characters and productivityrdquoPakistan Journal of Biological Sciences vol 4 no 1 pp 37ndash402001
[53] F R Piscitelli ldquoPrincipals caracteres exomorfologicos de cul-tures de cana de azucarrdquo TUC Iindustraly Agricos vol 7 pp 49ndash57 1994 (Spanish)
[54] A H D Brown and B S Weir ldquoMeasuring genetic variability inplant populationsrdquo in Isozyme in Plant Genetics Breeding S DTanksley and T J Orton Eds part A Elsevier Science 1983
[55] R Balakrishnan N V Nair and T V Sreenivasan ldquoA methodfor establishing a core collection of Saccharum officinarum Lgermplasm based on quantitative-morphological datardquo GeneticResources and Crop Evolution vol 47 no 1 pp 1ndash9 2000
[56] C A Nestor and B Q Reynaldo Sugarcane Variety Improve-ment in Southeast Asia and the Pacific for Enhanced andSustainable Productivity ndash Germplasm and Disease IndexingComponents Institute of Plant Breeding (IPB) College ofAgriculture (CA) UP Los Banos (UPLB) 2003
[57] A Maqbool J Muhammad and A Sagheer ldquoAgronomic traitsand morphological characteristics of some exotic varieties ofsugarcanerdquoPakistan Journal of Agricultural Research vol 19 no4 2006
over the regions for the 21 characters of the 211 sugarcaneaccessions studied and significant difference for the 6 regionsof origin for 6 characters (Table 3) The results suggestedthe occurrence of significant phenotypic variation betweenthe accessions as a whole Significant variations of differentsugarcane stalk and juice quality characters was also reportedin similar studies conducted elsewhere [26 27] Region-wise partitioning of the variance indicated significant within-region differences (p lt 005) among the populations withinalmost all regions for the characters NOI IL SH SD LLLW LA Brix Pol Purity SR MSCHA SCW CYHAand SY for 8 characters within Tigray for TC4MAP andTC5MAP within Oromiya and SNNPR Brix10MAP showedno significant difference within all regions
In general within-region variation was greater for stalkdiameter single cane weight millable stalk count and caneand sugar yield than for other characters for all the regionsAssuming that a significant portion of the phenotypic vari-ation is genetic it would be possible to make selection forany of this group of characters within a particular regionIt was apparent that the variance between accessions pooledover regions was greater than between regions Therefore inorder to sample the variation effectively it would be necessaryto sample the variable populations from different localitiesin a region The study in [28] on genetic diversity amongmain land USA sugarcane cultivars showed high geneticdiversitywithin populations of different regions than betweenpopulations
Duncanrsquos multiple range testing for regional means for allthe characters is shown in Table 4 Relatively much differenti-ationwas observed for stemdiameter and juice purity percentcompared to other parameters Important yield componentslike MSCHA SCW SH and SD have shown higher valuesin Tigray SNNPR and Oromiya regions High cane andsugar yield were recorded for accessions in Tigray regionfollowed by SNNPR Oromiya and introduction which werestatistically at par Statistically similar lower values of caneand sugar yield were recorded in Amhara and Benshangulcollections
Higher value of sucrose percent in cane was also observedfor accessions from Tigray and the lowest value was inAmhara accessions while other regions were statisticallysimilar Generally for most of the characters high valuewas obtained from accessions in Tigray region followed bySNNPROromiya and introduction Therefore the materialsfrom these regions can be used for selection of cane and sugaryield per se and in breeding program The local accessions inmany of the regions showed superiority over the introducedcommercial varieties currently in production for most of thecharacters including cane and sugar yield (Table 4)This maybe due to the adaptation and coevolution of the materialswith different stresses for long period of time in the countryrsquosagroecology These locally adapted genes conferring yieldadvantage could be harnessed through strategic crossing andselection for improvement of the crop Superiority of locallandraces over introduced varieties was reported [29 30]
The range of variation of the accession means establishedwide variation between the regions and the accessions withinthe regions for the characters studied (Table 5) Accordingly
the maximum score was 41 times the minimum for sproutcount one month after planting 32 for sprout count twomonths after planting 25 for tiller count four months afterplanting 22 for sugar yield 20 for cane yield 12 for millablestalk count 8 for stalk count 10 months after planting 7for single cane weight 6 for tiller count five months afterplanting and 2 for other yield components and sugar qual-ity parameters except hand refractometer brix reading tenmonths after planting and juice purity percent which scored1 The same trend was observed between accessions withina particular region However accessions within AmharaBenshangul-Gumz and introduction showed wider ranges ofvariation than those accessions within other regions for themajority of the characters The wide range of geographic andclimatic features of the regions in Ethiopia and the originaldifferentiation and further farmers selection for productionniches and uses for hundreds of years may have resultedin the accessions to possess a tremendously high degree ofmorphological variation The reason for introduced varietiesshowing high variation also could be due to broader geo-graphic spectrum fromwhere theywere initially acquired anddifference in their parental source and genealogical historyIt was demonstrated even with these limited numbers ofsamples (15 genotypes) that introduction of genetic materialshas practically broadened the genetic base of the sugarcanegermplasm for the breeding program
Coefficient of variation measured as the ratio of standarddeviation to the corresponding overall mean and expressedas percentage is useful to compare different charactersmeasured in different units It is also useful to compare samecharacter in different groups of populations with differentsample size mean and variance or different characters indifferent populations In the current study high coefficientsof variation were observed between regions and within eachregion for sprout as well as tiller counts in the monthsthe data was recorded stalk count 10 months after plant-ing and millable stalk count at harvest cane and sugaryield single cane weight and leaf area (Table 6) Thoughaccessions from a particular region were more variable fora specific character compared to other regions accessionsfromAmhara Benshangul-Gumz introduction and SNNPRwere more variable than accessions from other regions Thisresult demonstrated the tremendous variability of sugar-cane germplasm from these regions The accessions fromTigray and Oromiya had low coefficients of variation formany characters indicating relatively high within-regionuniformity The high coefficients of variation observed formost of the characters agreed well with those reportedin [31] for indigenous sugarcane of Brazil Similar resultswere also reported in sorghum [32] The different levels ofregional variability of a particular character could be due todifferences in forces of selection andor differences in theintensity of a particular selecting force It could also be dueto random effect and reduced sample parents of parentalmaterials moved differently by various human populationmigration and the interaction of these with Darwinianforces
Results of this study suggested the presence of wide rangeof variations of characters determined by univariate statistics
This is in agreement with previous studies on sugarcanegermplasm [33ndash35]
This is the first study to report information on geo-graphical pattern of variation in Ethiopian local sugarcanegermplasm which has been lacking so far Our results pro-vided experimental evidence on occurrence of geographicalvariation and significant within-region variation where it washigh in the regions such as Amhara Benshangul-Gumz andSNNPR
The overall patterns of similarity or difference betweenregions seemed to depend on environmental factors such asrainfall temperature length of growing season and altitudeWide variability of millable stalk count at harvest singlecane weight and plant height was observed among regionswhere these characters demonstrated high correlation withcane and sugar yield (Table 7) Therefore breeders can utilizeaccessions of regions showing variability for these charactersin selection programs and to design breeding strategies toproduce varieties with best commercial merits
Bivariate Statistics Correlation coefficients worked out onthe 21 quantitative characters are shown for the entire data(Table 7) between regions (Table 8) and within regions(data not shown) Results of the association studies for theentire data showed that cane and sugar yield had highlysignificant positive correlation with most of the quantitativestalk characters and juice quality parameters However theassociation of tiller counts 4 and 5 months after plantingstalk count 10 months after planting millable stalk countat harvest single cane weight and plant height with caneand sugar yield was stronger suggesting these componentcharacters asmain contributing factor to cane and sugar yieldIn fact sugar yield is the product of cane yield and sucrose per-cent cane surely being influenced with components affectingcane yield Similar trend was reported by earlier sugarcaneworkers [36 37] In sugarcane the cane and sugar yields areconsidered to be the complex characters The information onthe phenotypic interrelationship of cane yield and sugar yieldwith their component character per se would be of immensehelp to sugarcane breeders
Stem diameter had significant positive association withboth cane and sugar yield indicating the significance ofthis trait in improving both cane and sugar yields Similarreports have beenmade by earlier sugarcane workers [37 38]Stalk diameter was also positively and significantly correlatedwith plant height It is one of the traits that are related tolodging resistance [39] and the positive associations withheight would help in reducing the chance of lodging as heightincreases Among the physiological attributes included in thestudy leaf area had highly significant positive associationwith cane yield Similar observation was also made in [37]Sucrose percent which is an important juice quality attributehad highly significant positive association with sugar yieldand simultaneously cane yield also had highly significantassociation with sugar yield Based on the magnitude ofcorrelation coefficient values it can be said that cane yield ismuch more important than sucrose percent in determiningthe sugar yield and the same has also been reported by othersugarcane workers [37 40] Similar to observations made in
[37 41] results of the current study showed that all the qualityparameters like sucrose percent pol percent brix percentand purity percent were significantly correlated in positivedirection This indicated any of these juice quality traitscould be considered for selection leading to the simultaneousimprovement in the remaining quality traits
Ten of the 21 characters also showed significant positivecorrelations with altitude of the collection sites (Table 7) Asdescribed in [42] ecological characteristics have influencedthe genotypic constitution of landraces during domesticationand hence a relationship exists between the agroecology inthe collection site and the morphological characteristics ofthe landraces Thus positive correlation between collectionsite variables and plant characteristics would imply that thevariation between accessionsmay be related to agroecologicalvariations among the collection sites [43]
The correlation coefficient between number of internodesand altitude was negative and significant indicating thatother environmental factors (other than altitude) andornonenvironmental factorsmight account for the variation forthis particular character Since temperature decreases withan increase of altitude in Ethiopia [44 45] it is more likelythat temperature has exerted strong selection pressure innumber of internodes in the genotypes Information on therelationships between environmental factors of the collect-ing sites and morphophysiological variation of germplasmcould enhance the understanding of evolutionary adaptivepatterns which could assist germplasm collectors and users[46]
Character associations between region and within regionalso followed similar fashion like that of the entire dataNumber of sprout tiller and stalk counts and sugar qualityparameters showed significant positive correlations with caneand sugar yield
Though not significant and strong the correlation of leaflength and width and leaf area (leaf area = leaf length Xleaf width X 075) with cane and sugar yield was negativein the interregion correlations This kind of relationship wasalso exhibited between these leaf characteristics and yieldcomponents like tiller and stalks counts plant height andsingle cane weight and stalk diameter and sugar qualityparameters This may have been attributed to the variationof temperature with altitude and other environmental factorsacross the regions resulting in such type of relationshipSuch relationship also calls for the need to have enough dataon environmental variables of the collection sites such astemperature and rainfall if germplasm collection is requiredtomeet its objectives effectively and efficiently Unfortunatelydata on such useful environmental variables like temperatureand rainfall are lacking in the passport data of the currentEthiopian sugarcane germplasm collections The significanceof environmental factors of the collection sites as veryimportant determinants in structuring morphophysiolog-ical variations was reported in tetraploid wheat [22 4748]
Association of characters among yield its componentsand other economic traits is important for the interpretationof the patterns of variation and making selection in breed-ing program and combining several desirable attributes It
suggests the advantage of a scheme of selection for morethan one character at a time The correlation between char-acters may arise from linkage or from developmental geneticinteractions with orwithout a purely phenotypic componentIt could arise also due to differential phenotypic plasticityof characters themselves Within the limit of experimentalerror and environmental effects high correlation coefficientsbetween characters may show that the characters sharesome common element of genetic control (ie pleiotropylinkage) between genes or else from independently controlledcharacters responding similarly to geographic variation inselection pressures [49] The interregion correlation coef-ficient between the characters measures the consistency oftheir patterns of regional variation while the intraregioncorrelation coefficient measures the association arising fromgenetic factors but is not affected by regional variation[50] Since this study showed significant positive correlationsintraregionally for some character combinations it wouldseem that common genetic control might be playing a role inbringing about correlations between the various charactersIt appeared that different response to regional variationwas playing a greater role than different genetic controlas shown by the many more significant and moderate tohigh correlation coefficients intraregionally than interregion-ally
322 Qualitative Characters The frequency distribution forthe 16 qualitative characters of germplasm samples by regionsin Ethiopia and of the exotic accessions is shown in Supple-mental Table 5 For the purpose of avoiding redundancy andimproving readability the discrete characters are presented bygrouping related traits together
BudCushion andRelativeDegree of BudExtension Evaluationof the presence and absence of bud cushion revealed thatgreat proportion (70) of sugarcane landraces from Ethiopiado not have bud cushion Majority of the sample germplasmin all regions belong to this phenotypic class Approximatelyone-half (53) of the collection have their bud extendedabove the growth ring followed by those (41) having budtouching the growth ring
Relative Bud Shape Among the phenotypic classes for thischaracter tall deltoid was dominant (31) followed by ovate(17) and narrow ovate (12) While local accessions inmost regions had tall deltoid bud shape 50 of the intro-duced accessions belong to two phenotypic classes roundwith central germpore (30) and 20 having pentagonalbud shape Equal proportions (19) of the samples fromBenshangul-Gumz were Ovate and Ovate with emarginatebasal wing
Canopy Structure Relative Plant Erectness Colour of theLeaves and Colour of the Exposed Rind Out of the eightphenotypic classes observed compact tip droopy was themost frequent (35) Open semidroopy and open droopyclasses had fairly equal distribution With regard to relativeplant erectness seventy-five percent of the germplasm werefound to fall in twophenotypic classes erect (41) and almost
erect (34) for the whole region and country Most of theaccessions (90) showed light green or green leaf colourbut 20 from introduced collections were greenish yellowAmong phenotypic classes for colour of the exposed rindbrownish yellow yellow brown and yellowish green were themost frequent with fairly equal distribution
Relative Shape of Dewlap Double crescent had larger fre-quency (22) while squarish descending and flaring typeof dewlap shape had fairly equal proportions Seventeenpercent of germplasm from Tigray region had squarishdeltoid dewlap shape
Relative Degree of Internode Alignment and Internode ShapeForty-four percent of the sample germplasm were slightlyzigzag while there was almost equal frequency for straightand zigzag internode alignment for the entire region andcountry Straight or nearly straight (slightly zigzag) internodealignment of cane stalks is a very important character formechanized farming and postharvest handling in sugarcane[51] Forty percent of the collections were observed havingconcave-convex type of internode shape while 29 wascylindrical and 14 conoidal Other classes had fairly equalproportion
Type of Auricle and Relative Shape of Ligule An equalproportion of the germplasm totally constituting 84 ofthe collection belong to four phenotypic classes namelythose with no auricle and others with transitional and shortand lanceolate auricles Sixteen percent of the collectionsalso showed equal frequency for the remaining classesBenshangul-Gumz and Tigray regions had each equal 13accessions with falcate type of auricle Sixty-seven percentof the collections belong to three ligule shape classes cres-cent with lozenge (27) broad-crescent (21) and linear-crescent (19) The germplasm had almost equal proportionfor the other phenotypic classes of this character Nineteenpercent of the accessions fromBenshangul-Gumz region haddeltoid shape of ligule
Stalk Corky Cracks Corky Patches Growth Cracks and BudGroove Eighty-nine percent of the germplasm across regionshad no stalk corky cracks with only eleven percent exhibitingthis character All the germplasm in Gambella Harari andSomali had no stalk corky cracks On the other hand eighty-one percent of the germplasm for the whole region showedthe presence of stalk corky patches All the local germplasmin Benshangul-Gumz Gambella and Tigray had stalk corkypatches whereas 75 of the germplasm from Harari Region67 from Somali Region and 24 from Oromia Regionhad no stalk corky patches In the entire region 75 ofthe germplasm had no stalk growth cracks with only 25showing this character Sixty-three percent of the germplasmwere found to have bud groove Howevermost of the samplesin Harari (75) Amhara (63) and Oromia (53) Regionshad no bud groove
The results of this study indicated the wide distributionof phenotypic classes for the characters considered whichindicates the existence of different races and combination
Advances in Agriculture 15
of races in local genotypes of different regions in EthiopiaThis diversity of germplasmbased onphenotypicmarkers canhelp sugarcane breeders in identifying the identity of geno-types as well as maintaining genetic diversity Differencesin morphological characters of different sugarcane varietieshave been reported [52] Piscitelli [53] showed importantexomorphological qualitative characters of sugarcane varietythat are not influenced by environmental factors and thus canbe used as a selection tool in any breeding program
The amount of phenotypic diversity estimates based onthe ShannonndashWeaver diversity index (H1015840) are shown inTable 9 The 16 characters differed in their distribution aswell as the amount of variation Individual traits showed adifferent pattern of variation among accessions Estimatesof H1015840 for individual characters varied from 049 for stalkcorky cracks to 100 for bud groove with an overall meanof 080 Most of the characters were highly polymorphicwhereas relative plant erectness and presence or absenceof stalk corky patches scored moderate values The highdiversity values for the characters showed a wide variabilityamong genotypes According to [54] this index is also usedin genetic resource studies as a convenient measure of bothallelic richness and allelic evenness when using geneticdata
The highest diversity computed was on presence orabsence of bud groove with a diversity index of 100indicating that this character is very variable in the localsugarcane germplasm On the other hand the descriptorcomputed with the lowest diversity index value (049) wasthe presence or absence of stalk corky cracks indicatingthat this character is selected in the development of thegenotypes In agreement with the present study [55] workingon S officinarum accessions from the world collection ofsugarcane germplasm obtained a low index of 025 and 035for presence or absence of stalk corky patches and cracksrespectively
Their report also showed high diversity index values forinternode alignment growth cracks internode shape budcushion bud extension bud groove and ligule shape Sim-ilarly [56] experimenting on field collections of sugarcaneaccessions in Philippine reported high ShannonndashWeaverdiversity for stalk growth cracks bud cushion stalk corkypatch stalk corky cracks bud extension and auricle outershape Unlike the present study they have reported high indexfor stalk corky cracksMedium index values were reported forstalk internode shape and low index values for ligule shapeand bud shape High variability of sugarcane varieties fordewlap shape was also reported in [57]
The H1015840 pooled across characters by region ranged from079 in Oromiya to 085 in Tigray with an overall aver-age of 081 (Table 8) Most of the regions showed highShannonndashWeaver diversity index The populations that hadthe highest H1015840 were that of Tigray and introduction The H1015840averaged over regions for different characters was found torange from 044 for presence or absence of stalk corky cracksto 094 for relative degree of internode alignment and budgroove with an overall average of 081 (Table 9)
Assessment of the ShannonndashWeaver index also showedsensitivity to both the number of descriptor classes and the
distribution within phenotypic classes in each region Amongregions the mean values for the ShannonndashWeaver index arenot significantly different but differences were found whenindividual characters were considered These observationssuggest that some regions have high diversity for particulartraits while in other regions selective pressures might havereduced the variation to certain genotypes
4 Conclusions
The large number of landraces observed during the cur-rent study demonstrates the existence of diverse geneticresources of sugarcane in Ethiopia The landraces wouldserve to broaden the genetic base of the available sugarcanegermplasm Generally distribution of the overall phenotypicdiversity among sugarcane landraces in Ethiopia is unevenOur results showed that there was a wide range of variationthat existed in the local sugarcane landraces studied both atregional and within region levels Selection and differenti-ation of types might have taken place along a geographicalpattern perhaps associated with climate and use The presentstudy contributes to updating sugarcane descriptors adoptedfrom USDA-ARS as well as Bioversity passport data forthe future collection and evaluation As Ethiopian SugarCorporation is in the course of launching breeding programthe observed substantial variation of landraces would enablesugarcane breeders to design and practice breeding andselection programs to improve the crop
Data Availability
The data used to support the findings of this study areavailable from the corresponding author upon request
Conflicts of Interest
The authors declare that they have no conflicts of interest
Acknowledgments
The authors are grateful for the financial grant of SugarCorporation of Ethiopia Many thanks are due to EthiopianBiodiversity Institute (EBI) for its technical advice duringcollection of the local sugarcane genotypes Sincere thanksare forwarded to Ethiopian smallholder farmers who pre-served sugarcane germplasm and shared their knowledgeand experience Regional Zonal and District (ldquoWeredardquo)Agriculture Bureaus across Ethiopia are acknowledged fortheir support during the study
Supplementary Materials
These supplementary materials contain different tables sup-porting the results of the study These include passport dataof sugarcane germplasm collected and related informationon the materials local sugarcane germplasm collected andused formorphological diversity study introduced sugarcane
16 Advances in Agriculture
Table9Sh
anno
n-WeaverD
iversityIndices(H1015840)a
ndmeanandsta
ndarderror(plusmnS
E)of
land
racesineach
region
basedon
16discretecharacters
Characters
Amha
raBe
nsha
ngul-G
umz
Introd
uced
Oromia
SNNP
Tigray
ELb
Total
Presence
orabsenceo
fbud
cushion(BUDCUSH
ION)
083
095
095
089
082
098
090
088
Relativ
edegreeo
fbud
extension(BUDEX
TEND)
077
079
093
074
072
087
080
078
Relativ
ebud
Shape(BU
DSH
APE
)078
096
089
081
085
078
084
078
Canop
ystructure(C
ANOPY
)089
098
095
081
078
096
089
084
Relativ
eshape
ofdewlap(D
EWLA
PSHAP)
087
094
092
090
082
096
090
086
Relativ
eplant
erectness(ER
ECT)
080
082
050
067
071
088
073
064
Relativ
edegreeo
finterno
dealignm
ent(IN
ALIGN)
096
076
098
097
098
100
094
098
Relativ
einterno
deshape(IN
SHAPE
)086
069
095
077
083
091
083
082
Color
oftheleaves(LE
AFC
OLO
R)087
071
090
064
074
071
076
076
Type
ofou
tera
uricle(A
URI
CLE
OUT)
080
092
087
079
083
093
086
082
Relativ
eshape
ofligule(LIGSH
APE
)086
094
082
085
090
083
087
083
Presence
orabsenceo
fstalkcorkyc
racks(ST
ALK
CORK
C)
044
054
035
055
051
025
044
049
Presence
orabsenceo
fstalkcorkyp
atches(STA
LKCORK
P)066
000
072
079
059
087
060
069
Presence
orabsenceo
fstalkgrow
thcracks
(STA
LKCRA
CK)
066
100
088
062
088
099
084
080
Color
ofthee
xposed
rind(R
INDCOLE
)085
096
089
084
086
076
086
085
Presence
orabsenceo
fbud
groo
ve(BUDGRO
OVE)
095
100
072
100
100
099
094
100
Means
080plusmn0
03
081plusmn0
06
083plusmn0
04
079plusmn0
03
080plusmn0
03
085plusmn0
05
081plusmn0
03
080
b EL=
averageo
fentire
locatio
n
Advances in Agriculture 17
varieties used for morphological diversity study and fre-quency distribution for 16 qualitative characters in sugarcaneby regions in Ethiopia and country of sources of introducedgenotypes (Supplementary Materials)
References
[1] ldquoSpreading the word about clean solutions from sugarcanrdquoAccessed 04072018 httpwwwsugarcaneorg
[2] Central Statistics Agency (CSA) ldquoAgricultural sample surveyArea and production of major crops private peasant holdingsrdquoStatistical Bulletin Addis Ababa Ethiopia 2017
[3] T Esayas Exploration and Collection Characterization GeneticDiversity Analysis and Association of Traits for Yield and YieldComponents of Sugarcane (Saccharumspp) in Ethiopia [MasterThesis] Haramaya University Dire Dawa Ethiopia 2014
[4] J R Harlan Crops and Man American Society of AgronomyCrop Science Society of America Madison WI USA 2ndedition 1992
[5] R W Allard Principles of Plant Breeding JohnWilley and SonsNew York NY USA 2nd edition 1999
[6] B T Roach and J Daniels ldquoA review of the origin and improve-ment of sugarcanerdquo in Proceedings of Coper sugar InternationalSugarcane Workshop pp 1ndash32 1987
[7] P P Patil S S Patil and D U Patel ldquoGenetic diversity in sug-arcane (Saccharum spp Complex)rdquo Journal of Pharmacognosyand Phytochemistry vol 6 no 6 pp 570ndash573 2017
[8] R B Singh B Singh andR K Singh ldquoStudy of genetic diversityof sugarcane (Saccharum) species and commercial varietiesthrough TRAP molecular markersrdquo Indian Journal of PlantPhysiology vol 22 no 3 pp 332ndash338 2017
[9] R W Allard ldquoPopulation structure and sampling methodsrdquo inGenetic Resources in PlantsmdashTheir Exploration and Conserva-tion O H Frankel and E Bennett Eds IBP Handbook no 11pp 97ndash107 FA Davis Company Philadelphia PA USA 1970
[10] D R Marshall and A H D Brown ldquoOptimum samplingstrategies in genetic conservationrdquo inCrop Genetic Resources forToday and Tomorrow O H Frankel and J G Hawkes Eds pp53ndash80 Cambridge University Press 1975
[11] J G Hawkes Crop Genetic Resources Field Collection ManualPudoc Wagnningen Netherland 1980
[12] J G Hawkes The Diversity of Crop Plants Harvard UniversityPress London England 1983
[13] B P K Yerima Manual for Good Laboratory Practice Foodand Agriculture Organization of United Nation Addis AbabaEthiopia 1993
[14] A Walkley and I A Black ldquoAn examination of the degtjareffmethod for determining soil organic matter and a proposedmodification of the chromic acid titrationmethodrdquo Soil Sciencevol 37 no 1 pp 29ndash38 1934
[15] S R Olsen and L E Sommers ldquoPhosphorusrdquo in AgronomyMonograph A L Page et al Ed Methods of soil analysis part2 pp 403ndash430 ASA and SSSAMadisonWI USA 2nd edition1982
[16] R H Bray and L T Kurtz ldquoDetermination of total organic andavailable forms of phosphorus in soilsrdquo Soil Science vol 59 no1 pp 39ndash46 1945
[17] S Sahlemedhin andB Taye ldquoProcedures for soil and plant anal-ysisrdquo Tech Rep 74 National Soil Research Center EthiopianAgricultural Research Organization 2000
[18] F C Stickler S Weaden and AW Pauli ldquoLeaf area determina-tion in grain sorghumrdquo Journal of Agronomy and Crop Sciencevol 53 pp 187-188 1961
[19] Munsell Color ChartThe Munsell Book of Color Matte Collec-tion Munsell Color Chart NewWindsor NY USA 1990
[20] H O Hartley ldquoThe maximum F-ratio as a short cut test forheterogeneity of variancesrdquo Biometrika vol 37 pp 308ndash3121950
[21] L Pecetti P Annicchiarico and A B Damania ldquoBiodiversity ina germplasm collection of durumwheatrdquo Euphytica vol 60 no3 pp 229ndash238 1992
[22] L Pecetti and A B Damania ldquoGeographic variation intetraploid wheat (Triticum turgidum ssp turgidum convardurum) landraces from two provinces in Ethiopiardquo GeneticResources and Crop Evolution vol 43 no 5 pp 395ndash407 1996
[23] S K Jain C O Qualset G M Bhatt and K K Wu ldquoGeo-graphical patterns of phenotypic diversity in a world collectionof durumwheatsrdquoCrop Science vol 15 no 5 pp 700ndash704 1975
[24] K K Nkongolo L Chinthu M Malusi and Z Vokhiwa ldquoPar-ticipatory variety selection and characterization of sorghum(Sorghum bicolour (L) Moench) elite accessions from Malaw-ian gene pool using farmer and breeder knowledgerdquo AfricanJournal of Agricultural Research vol 3 pp 273ndash283 2008
[25] M B Semira ldquoInnovative methods and metrics for agricul-ture and nutrition actions soil quality and human nutritionin Ethiopiardquo Leverhulme Center for Integrative Research onAgriculture and Health (LCIRAH) 2017
[26] I A Khan A Khatri M A Siddiqui G S Nizamani and SRaza ldquoPerformance of promising sugarcane clone for yield andquality traits in different ecological zones of Sindhirdquo PakistanJournal of Botany vol 36 no 1 pp 83ndash92 2004
[27] M F Perera M E Arias D Costilla et al ldquoGenetic diversityassessment and genotype identification in sugarcane based onDNAmarkers andmorphological traitsrdquo Euphytica vol 185 no3 pp 491ndash510 2012
[28] C G Neil M Kay and C C Jack ldquoDiversity among mainlandUSA sugarcane cultivars examined by SSR genotypingrdquo Journalof the American Society of Sugar Cane Technologists vol 29 pp36ndash52 2009
[29] S SchenckMWCrepeauK KWu P HMooreQ Yu andRMing ldquoGenetic diversity and relationships in native HawaiianSaccharum officinarum sugarcanerdquo Journal of Heredity vol 95no 4 pp 327ndash331 2004
[30] R Gebretsadik H Shimelis M D Laing P Tongoona andN Mandefro ldquoA diagnostic appraisal of the sorghum farmingsystem and breeding priorities in Striga infested agro-ecologiesof Ethiopiardquo Agricultural Systems vol 123 pp 54ndash61 2014
[31] L E V Cesar A T Bruzi J A R Nunes et al ldquoGeneticdivergence of sugarcane varieties collected in the region ofLavras Minas Gerais Brazilrdquo Genetics and Molecular Researchvol 14 no 4 pp 14043ndash14049 2015
[32] C Grenier P J Bramel J A Dahlberg et al ldquoSorghums of theSudan analysis of regional diversity and distributionrdquo GeneticResources and Crop Evolution vol 51 no 5 pp 489ndash500 2004
[33] N V Nair R Balakrishnan and T V Sreenivasan ldquoVariabilityfor quantitative traits in exotic hybrid germplasm of sugarcanerdquoGenetic Resources and Crop Evolution vol 45 no 5 pp 459ndash464 1998
[34] R Nagarajan S Alarmelu and R M Shanthi ldquoStudies onvariation in interspecific hybrids of Saccharumrdquo Sugar Tech vol2 no 3 pp 42ndash46 2000
18 Advances in Agriculture
[35] D N Kamat and J R P Singh ldquoVariability in Sugarcane underrain fed conditionrdquo Sugar Technology Short Communicationsvol 3 no 1-2 pp 65ndash67 2001
[36] B S Rishipal A S M Chaudhary and S P Kadian ldquoStudieson character interrelationship in clonal generation of sugarcane(Saccharum complex hybrids)rdquo Indian Sugar vol 47 no 11 pp907ndash911 1998
[37] S Thippeswamy S T Kajjidoni P M Salimath and J V GoudldquoCorrelation and path analysis for cane yield juice quality andtheir component traits in sugarcanerdquo Short Communication vol5 no 1-2 pp 65ndash72 2003
[38] P S Verma S Pal and N K Karma ldquoGenetic variability andcorrelation studies in sugarcanerdquo Indian Sugar vol 49 pp 125ndash128 1999
[39] H A Esechie J W Maranville and W M Ross ldquoRelationshipof stalk morphology and chemical composition to lodgingresistance in sorghumrdquo Crop Science vol 17 no 4 p 609 1977
[40] P K Das B C Jena and N Nayak ldquoCharacter association andpath analysis of sugar yield in sugarcanerdquo Indian Sugar vol 46no 10 pp 805ndash808 1997
[41] K J Sreekumar M Kuriakose M Thomas D Alexander andS Santhakumari ldquoVariability heritability and correlation onstudies on the yield and quality characters of sugarcanerdquo IndianSugar vol 44 no 4 pp 243ndash249 1994
[42] J R Harlan J M de Wet and E G Price ldquoComparativeevolution of cerealsrdquo Evolution vol 27 no 2 pp 311ndash325 1973
[43] A Elings ldquoDurum wheat landraces from syria II patterns ofvariationrdquo Euphytica vol 54 no 3 pp 231ndash243 1991
[44] K Tato ldquoRainfall in Ethiopiardquo Ethiopian Geographical Journalvol 2 pp 28ndash36 1964
[45] A B L Stemler J R Harlan and J M J deWet ldquoThe sorghumsof ethiopiardquo Economic Botany vol 31 no 4 pp 446ndash460 1977
[46] P Annicchiarico L Pecetti and A B Damania ldquoRelationshipsbetween phenotypic variation and climatic factors at collectingsites in durum wheat landracesrdquo Hereditas vol 122 no 2 pp163ndash167 1995
[47] A B Damania L Pecetti C O Qualset and B O HumeidldquoDiversity and geographic distribution of adaptive traits inTriticum turgidum L (durum group) wheat landraces fromTurkeyrdquoGenetic Resources and Crop Evolution vol 43 no 5 pp409ndash422 1996
[48] A Ayana and E Bekele ldquoGeographical patterns of morpho-logical variation in sorghum (Sorghum bicolor (L) Moench)germplasm from Ethiopia and Eritrea quantitative charactersrdquoEuphytica vol 115 no 2 pp 91ndash104 2000
[49] D S Falconer Introduction to Quantitative Genetics JohnWilley and Sons Inc New York NY USA 3rd edition 1993
[50] R S Thorpe ldquoBiochemical analysis of geographical variationand racial affinitiesrdquo Biology Review vol 51 pp 407ndash452 1976
[51] M Farooq ldquoMorphological and agricultural characteristicsof sugarcane in relation to mechanized agriculture at NARCIslamabadrdquo Portu Rico Journal vol 1 pp 51ndash61 1989
[52] M Akhtar N N Elahi and M Ashraf ldquoEvaluation of exoticsugarcane varieties for agronomic characters and productivityrdquoPakistan Journal of Biological Sciences vol 4 no 1 pp 37ndash402001
[53] F R Piscitelli ldquoPrincipals caracteres exomorfologicos de cul-tures de cana de azucarrdquo TUC Iindustraly Agricos vol 7 pp 49ndash57 1994 (Spanish)
[54] A H D Brown and B S Weir ldquoMeasuring genetic variability inplant populationsrdquo in Isozyme in Plant Genetics Breeding S DTanksley and T J Orton Eds part A Elsevier Science 1983
[55] R Balakrishnan N V Nair and T V Sreenivasan ldquoA methodfor establishing a core collection of Saccharum officinarum Lgermplasm based on quantitative-morphological datardquo GeneticResources and Crop Evolution vol 47 no 1 pp 1ndash9 2000
[56] C A Nestor and B Q Reynaldo Sugarcane Variety Improve-ment in Southeast Asia and the Pacific for Enhanced andSustainable Productivity ndash Germplasm and Disease IndexingComponents Institute of Plant Breeding (IPB) College ofAgriculture (CA) UP Los Banos (UPLB) 2003
[57] A Maqbool J Muhammad and A Sagheer ldquoAgronomic traitsand morphological characteristics of some exotic varieties ofsugarcanerdquoPakistan Journal of Agricultural Research vol 19 no4 2006
This is in agreement with previous studies on sugarcanegermplasm [33ndash35]
This is the first study to report information on geo-graphical pattern of variation in Ethiopian local sugarcanegermplasm which has been lacking so far Our results pro-vided experimental evidence on occurrence of geographicalvariation and significant within-region variation where it washigh in the regions such as Amhara Benshangul-Gumz andSNNPR
The overall patterns of similarity or difference betweenregions seemed to depend on environmental factors such asrainfall temperature length of growing season and altitudeWide variability of millable stalk count at harvest singlecane weight and plant height was observed among regionswhere these characters demonstrated high correlation withcane and sugar yield (Table 7) Therefore breeders can utilizeaccessions of regions showing variability for these charactersin selection programs and to design breeding strategies toproduce varieties with best commercial merits
Bivariate Statistics Correlation coefficients worked out onthe 21 quantitative characters are shown for the entire data(Table 7) between regions (Table 8) and within regions(data not shown) Results of the association studies for theentire data showed that cane and sugar yield had highlysignificant positive correlation with most of the quantitativestalk characters and juice quality parameters However theassociation of tiller counts 4 and 5 months after plantingstalk count 10 months after planting millable stalk countat harvest single cane weight and plant height with caneand sugar yield was stronger suggesting these componentcharacters asmain contributing factor to cane and sugar yieldIn fact sugar yield is the product of cane yield and sucrose per-cent cane surely being influenced with components affectingcane yield Similar trend was reported by earlier sugarcaneworkers [36 37] In sugarcane the cane and sugar yields areconsidered to be the complex characters The information onthe phenotypic interrelationship of cane yield and sugar yieldwith their component character per se would be of immensehelp to sugarcane breeders
Stem diameter had significant positive association withboth cane and sugar yield indicating the significance ofthis trait in improving both cane and sugar yields Similarreports have beenmade by earlier sugarcane workers [37 38]Stalk diameter was also positively and significantly correlatedwith plant height It is one of the traits that are related tolodging resistance [39] and the positive associations withheight would help in reducing the chance of lodging as heightincreases Among the physiological attributes included in thestudy leaf area had highly significant positive associationwith cane yield Similar observation was also made in [37]Sucrose percent which is an important juice quality attributehad highly significant positive association with sugar yieldand simultaneously cane yield also had highly significantassociation with sugar yield Based on the magnitude ofcorrelation coefficient values it can be said that cane yield ismuch more important than sucrose percent in determiningthe sugar yield and the same has also been reported by othersugarcane workers [37 40] Similar to observations made in
[37 41] results of the current study showed that all the qualityparameters like sucrose percent pol percent brix percentand purity percent were significantly correlated in positivedirection This indicated any of these juice quality traitscould be considered for selection leading to the simultaneousimprovement in the remaining quality traits
Ten of the 21 characters also showed significant positivecorrelations with altitude of the collection sites (Table 7) Asdescribed in [42] ecological characteristics have influencedthe genotypic constitution of landraces during domesticationand hence a relationship exists between the agroecology inthe collection site and the morphological characteristics ofthe landraces Thus positive correlation between collectionsite variables and plant characteristics would imply that thevariation between accessionsmay be related to agroecologicalvariations among the collection sites [43]
The correlation coefficient between number of internodesand altitude was negative and significant indicating thatother environmental factors (other than altitude) andornonenvironmental factorsmight account for the variation forthis particular character Since temperature decreases withan increase of altitude in Ethiopia [44 45] it is more likelythat temperature has exerted strong selection pressure innumber of internodes in the genotypes Information on therelationships between environmental factors of the collect-ing sites and morphophysiological variation of germplasmcould enhance the understanding of evolutionary adaptivepatterns which could assist germplasm collectors and users[46]
Character associations between region and within regionalso followed similar fashion like that of the entire dataNumber of sprout tiller and stalk counts and sugar qualityparameters showed significant positive correlations with caneand sugar yield
Though not significant and strong the correlation of leaflength and width and leaf area (leaf area = leaf length Xleaf width X 075) with cane and sugar yield was negativein the interregion correlations This kind of relationship wasalso exhibited between these leaf characteristics and yieldcomponents like tiller and stalks counts plant height andsingle cane weight and stalk diameter and sugar qualityparameters This may have been attributed to the variationof temperature with altitude and other environmental factorsacross the regions resulting in such type of relationshipSuch relationship also calls for the need to have enough dataon environmental variables of the collection sites such astemperature and rainfall if germplasm collection is requiredtomeet its objectives effectively and efficiently Unfortunatelydata on such useful environmental variables like temperatureand rainfall are lacking in the passport data of the currentEthiopian sugarcane germplasm collections The significanceof environmental factors of the collection sites as veryimportant determinants in structuring morphophysiolog-ical variations was reported in tetraploid wheat [22 4748]
Association of characters among yield its componentsand other economic traits is important for the interpretationof the patterns of variation and making selection in breed-ing program and combining several desirable attributes It
suggests the advantage of a scheme of selection for morethan one character at a time The correlation between char-acters may arise from linkage or from developmental geneticinteractions with orwithout a purely phenotypic componentIt could arise also due to differential phenotypic plasticityof characters themselves Within the limit of experimentalerror and environmental effects high correlation coefficientsbetween characters may show that the characters sharesome common element of genetic control (ie pleiotropylinkage) between genes or else from independently controlledcharacters responding similarly to geographic variation inselection pressures [49] The interregion correlation coef-ficient between the characters measures the consistency oftheir patterns of regional variation while the intraregioncorrelation coefficient measures the association arising fromgenetic factors but is not affected by regional variation[50] Since this study showed significant positive correlationsintraregionally for some character combinations it wouldseem that common genetic control might be playing a role inbringing about correlations between the various charactersIt appeared that different response to regional variationwas playing a greater role than different genetic controlas shown by the many more significant and moderate tohigh correlation coefficients intraregionally than interregion-ally
322 Qualitative Characters The frequency distribution forthe 16 qualitative characters of germplasm samples by regionsin Ethiopia and of the exotic accessions is shown in Supple-mental Table 5 For the purpose of avoiding redundancy andimproving readability the discrete characters are presented bygrouping related traits together
BudCushion andRelativeDegree of BudExtension Evaluationof the presence and absence of bud cushion revealed thatgreat proportion (70) of sugarcane landraces from Ethiopiado not have bud cushion Majority of the sample germplasmin all regions belong to this phenotypic class Approximatelyone-half (53) of the collection have their bud extendedabove the growth ring followed by those (41) having budtouching the growth ring
Relative Bud Shape Among the phenotypic classes for thischaracter tall deltoid was dominant (31) followed by ovate(17) and narrow ovate (12) While local accessions inmost regions had tall deltoid bud shape 50 of the intro-duced accessions belong to two phenotypic classes roundwith central germpore (30) and 20 having pentagonalbud shape Equal proportions (19) of the samples fromBenshangul-Gumz were Ovate and Ovate with emarginatebasal wing
Canopy Structure Relative Plant Erectness Colour of theLeaves and Colour of the Exposed Rind Out of the eightphenotypic classes observed compact tip droopy was themost frequent (35) Open semidroopy and open droopyclasses had fairly equal distribution With regard to relativeplant erectness seventy-five percent of the germplasm werefound to fall in twophenotypic classes erect (41) and almost
erect (34) for the whole region and country Most of theaccessions (90) showed light green or green leaf colourbut 20 from introduced collections were greenish yellowAmong phenotypic classes for colour of the exposed rindbrownish yellow yellow brown and yellowish green were themost frequent with fairly equal distribution
Relative Shape of Dewlap Double crescent had larger fre-quency (22) while squarish descending and flaring typeof dewlap shape had fairly equal proportions Seventeenpercent of germplasm from Tigray region had squarishdeltoid dewlap shape
Relative Degree of Internode Alignment and Internode ShapeForty-four percent of the sample germplasm were slightlyzigzag while there was almost equal frequency for straightand zigzag internode alignment for the entire region andcountry Straight or nearly straight (slightly zigzag) internodealignment of cane stalks is a very important character formechanized farming and postharvest handling in sugarcane[51] Forty percent of the collections were observed havingconcave-convex type of internode shape while 29 wascylindrical and 14 conoidal Other classes had fairly equalproportion
Type of Auricle and Relative Shape of Ligule An equalproportion of the germplasm totally constituting 84 ofthe collection belong to four phenotypic classes namelythose with no auricle and others with transitional and shortand lanceolate auricles Sixteen percent of the collectionsalso showed equal frequency for the remaining classesBenshangul-Gumz and Tigray regions had each equal 13accessions with falcate type of auricle Sixty-seven percentof the collections belong to three ligule shape classes cres-cent with lozenge (27) broad-crescent (21) and linear-crescent (19) The germplasm had almost equal proportionfor the other phenotypic classes of this character Nineteenpercent of the accessions fromBenshangul-Gumz region haddeltoid shape of ligule
Stalk Corky Cracks Corky Patches Growth Cracks and BudGroove Eighty-nine percent of the germplasm across regionshad no stalk corky cracks with only eleven percent exhibitingthis character All the germplasm in Gambella Harari andSomali had no stalk corky cracks On the other hand eighty-one percent of the germplasm for the whole region showedthe presence of stalk corky patches All the local germplasmin Benshangul-Gumz Gambella and Tigray had stalk corkypatches whereas 75 of the germplasm from Harari Region67 from Somali Region and 24 from Oromia Regionhad no stalk corky patches In the entire region 75 ofthe germplasm had no stalk growth cracks with only 25showing this character Sixty-three percent of the germplasmwere found to have bud groove Howevermost of the samplesin Harari (75) Amhara (63) and Oromia (53) Regionshad no bud groove
The results of this study indicated the wide distributionof phenotypic classes for the characters considered whichindicates the existence of different races and combination
Advances in Agriculture 15
of races in local genotypes of different regions in EthiopiaThis diversity of germplasmbased onphenotypicmarkers canhelp sugarcane breeders in identifying the identity of geno-types as well as maintaining genetic diversity Differencesin morphological characters of different sugarcane varietieshave been reported [52] Piscitelli [53] showed importantexomorphological qualitative characters of sugarcane varietythat are not influenced by environmental factors and thus canbe used as a selection tool in any breeding program
The amount of phenotypic diversity estimates based onthe ShannonndashWeaver diversity index (H1015840) are shown inTable 9 The 16 characters differed in their distribution aswell as the amount of variation Individual traits showed adifferent pattern of variation among accessions Estimatesof H1015840 for individual characters varied from 049 for stalkcorky cracks to 100 for bud groove with an overall meanof 080 Most of the characters were highly polymorphicwhereas relative plant erectness and presence or absenceof stalk corky patches scored moderate values The highdiversity values for the characters showed a wide variabilityamong genotypes According to [54] this index is also usedin genetic resource studies as a convenient measure of bothallelic richness and allelic evenness when using geneticdata
The highest diversity computed was on presence orabsence of bud groove with a diversity index of 100indicating that this character is very variable in the localsugarcane germplasm On the other hand the descriptorcomputed with the lowest diversity index value (049) wasthe presence or absence of stalk corky cracks indicatingthat this character is selected in the development of thegenotypes In agreement with the present study [55] workingon S officinarum accessions from the world collection ofsugarcane germplasm obtained a low index of 025 and 035for presence or absence of stalk corky patches and cracksrespectively
Their report also showed high diversity index values forinternode alignment growth cracks internode shape budcushion bud extension bud groove and ligule shape Sim-ilarly [56] experimenting on field collections of sugarcaneaccessions in Philippine reported high ShannonndashWeaverdiversity for stalk growth cracks bud cushion stalk corkypatch stalk corky cracks bud extension and auricle outershape Unlike the present study they have reported high indexfor stalk corky cracksMedium index values were reported forstalk internode shape and low index values for ligule shapeand bud shape High variability of sugarcane varieties fordewlap shape was also reported in [57]
The H1015840 pooled across characters by region ranged from079 in Oromiya to 085 in Tigray with an overall aver-age of 081 (Table 8) Most of the regions showed highShannonndashWeaver diversity index The populations that hadthe highest H1015840 were that of Tigray and introduction The H1015840averaged over regions for different characters was found torange from 044 for presence or absence of stalk corky cracksto 094 for relative degree of internode alignment and budgroove with an overall average of 081 (Table 9)
Assessment of the ShannonndashWeaver index also showedsensitivity to both the number of descriptor classes and the
distribution within phenotypic classes in each region Amongregions the mean values for the ShannonndashWeaver index arenot significantly different but differences were found whenindividual characters were considered These observationssuggest that some regions have high diversity for particulartraits while in other regions selective pressures might havereduced the variation to certain genotypes
4 Conclusions
The large number of landraces observed during the cur-rent study demonstrates the existence of diverse geneticresources of sugarcane in Ethiopia The landraces wouldserve to broaden the genetic base of the available sugarcanegermplasm Generally distribution of the overall phenotypicdiversity among sugarcane landraces in Ethiopia is unevenOur results showed that there was a wide range of variationthat existed in the local sugarcane landraces studied both atregional and within region levels Selection and differenti-ation of types might have taken place along a geographicalpattern perhaps associated with climate and use The presentstudy contributes to updating sugarcane descriptors adoptedfrom USDA-ARS as well as Bioversity passport data forthe future collection and evaluation As Ethiopian SugarCorporation is in the course of launching breeding programthe observed substantial variation of landraces would enablesugarcane breeders to design and practice breeding andselection programs to improve the crop
Data Availability
The data used to support the findings of this study areavailable from the corresponding author upon request
Conflicts of Interest
The authors declare that they have no conflicts of interest
Acknowledgments
The authors are grateful for the financial grant of SugarCorporation of Ethiopia Many thanks are due to EthiopianBiodiversity Institute (EBI) for its technical advice duringcollection of the local sugarcane genotypes Sincere thanksare forwarded to Ethiopian smallholder farmers who pre-served sugarcane germplasm and shared their knowledgeand experience Regional Zonal and District (ldquoWeredardquo)Agriculture Bureaus across Ethiopia are acknowledged fortheir support during the study
Supplementary Materials
These supplementary materials contain different tables sup-porting the results of the study These include passport dataof sugarcane germplasm collected and related informationon the materials local sugarcane germplasm collected andused formorphological diversity study introduced sugarcane
16 Advances in Agriculture
Table9Sh
anno
n-WeaverD
iversityIndices(H1015840)a
ndmeanandsta
ndarderror(plusmnS
E)of
land
racesineach
region
basedon
16discretecharacters
Characters
Amha
raBe
nsha
ngul-G
umz
Introd
uced
Oromia
SNNP
Tigray
ELb
Total
Presence
orabsenceo
fbud
cushion(BUDCUSH
ION)
083
095
095
089
082
098
090
088
Relativ
edegreeo
fbud
extension(BUDEX
TEND)
077
079
093
074
072
087
080
078
Relativ
ebud
Shape(BU
DSH
APE
)078
096
089
081
085
078
084
078
Canop
ystructure(C
ANOPY
)089
098
095
081
078
096
089
084
Relativ
eshape
ofdewlap(D
EWLA
PSHAP)
087
094
092
090
082
096
090
086
Relativ
eplant
erectness(ER
ECT)
080
082
050
067
071
088
073
064
Relativ
edegreeo
finterno
dealignm
ent(IN
ALIGN)
096
076
098
097
098
100
094
098
Relativ
einterno
deshape(IN
SHAPE
)086
069
095
077
083
091
083
082
Color
oftheleaves(LE
AFC
OLO
R)087
071
090
064
074
071
076
076
Type
ofou
tera
uricle(A
URI
CLE
OUT)
080
092
087
079
083
093
086
082
Relativ
eshape
ofligule(LIGSH
APE
)086
094
082
085
090
083
087
083
Presence
orabsenceo
fstalkcorkyc
racks(ST
ALK
CORK
C)
044
054
035
055
051
025
044
049
Presence
orabsenceo
fstalkcorkyp
atches(STA
LKCORK
P)066
000
072
079
059
087
060
069
Presence
orabsenceo
fstalkgrow
thcracks
(STA
LKCRA
CK)
066
100
088
062
088
099
084
080
Color
ofthee
xposed
rind(R
INDCOLE
)085
096
089
084
086
076
086
085
Presence
orabsenceo
fbud
groo
ve(BUDGRO
OVE)
095
100
072
100
100
099
094
100
Means
080plusmn0
03
081plusmn0
06
083plusmn0
04
079plusmn0
03
080plusmn0
03
085plusmn0
05
081plusmn0
03
080
b EL=
averageo
fentire
locatio
n
Advances in Agriculture 17
varieties used for morphological diversity study and fre-quency distribution for 16 qualitative characters in sugarcaneby regions in Ethiopia and country of sources of introducedgenotypes (Supplementary Materials)
References
[1] ldquoSpreading the word about clean solutions from sugarcanrdquoAccessed 04072018 httpwwwsugarcaneorg
[2] Central Statistics Agency (CSA) ldquoAgricultural sample surveyArea and production of major crops private peasant holdingsrdquoStatistical Bulletin Addis Ababa Ethiopia 2017
[3] T Esayas Exploration and Collection Characterization GeneticDiversity Analysis and Association of Traits for Yield and YieldComponents of Sugarcane (Saccharumspp) in Ethiopia [MasterThesis] Haramaya University Dire Dawa Ethiopia 2014
[4] J R Harlan Crops and Man American Society of AgronomyCrop Science Society of America Madison WI USA 2ndedition 1992
[5] R W Allard Principles of Plant Breeding JohnWilley and SonsNew York NY USA 2nd edition 1999
[6] B T Roach and J Daniels ldquoA review of the origin and improve-ment of sugarcanerdquo in Proceedings of Coper sugar InternationalSugarcane Workshop pp 1ndash32 1987
[7] P P Patil S S Patil and D U Patel ldquoGenetic diversity in sug-arcane (Saccharum spp Complex)rdquo Journal of Pharmacognosyand Phytochemistry vol 6 no 6 pp 570ndash573 2017
[8] R B Singh B Singh andR K Singh ldquoStudy of genetic diversityof sugarcane (Saccharum) species and commercial varietiesthrough TRAP molecular markersrdquo Indian Journal of PlantPhysiology vol 22 no 3 pp 332ndash338 2017
[9] R W Allard ldquoPopulation structure and sampling methodsrdquo inGenetic Resources in PlantsmdashTheir Exploration and Conserva-tion O H Frankel and E Bennett Eds IBP Handbook no 11pp 97ndash107 FA Davis Company Philadelphia PA USA 1970
[10] D R Marshall and A H D Brown ldquoOptimum samplingstrategies in genetic conservationrdquo inCrop Genetic Resources forToday and Tomorrow O H Frankel and J G Hawkes Eds pp53ndash80 Cambridge University Press 1975
[11] J G Hawkes Crop Genetic Resources Field Collection ManualPudoc Wagnningen Netherland 1980
[12] J G Hawkes The Diversity of Crop Plants Harvard UniversityPress London England 1983
[13] B P K Yerima Manual for Good Laboratory Practice Foodand Agriculture Organization of United Nation Addis AbabaEthiopia 1993
[14] A Walkley and I A Black ldquoAn examination of the degtjareffmethod for determining soil organic matter and a proposedmodification of the chromic acid titrationmethodrdquo Soil Sciencevol 37 no 1 pp 29ndash38 1934
[15] S R Olsen and L E Sommers ldquoPhosphorusrdquo in AgronomyMonograph A L Page et al Ed Methods of soil analysis part2 pp 403ndash430 ASA and SSSAMadisonWI USA 2nd edition1982
[16] R H Bray and L T Kurtz ldquoDetermination of total organic andavailable forms of phosphorus in soilsrdquo Soil Science vol 59 no1 pp 39ndash46 1945
[17] S Sahlemedhin andB Taye ldquoProcedures for soil and plant anal-ysisrdquo Tech Rep 74 National Soil Research Center EthiopianAgricultural Research Organization 2000
[18] F C Stickler S Weaden and AW Pauli ldquoLeaf area determina-tion in grain sorghumrdquo Journal of Agronomy and Crop Sciencevol 53 pp 187-188 1961
[19] Munsell Color ChartThe Munsell Book of Color Matte Collec-tion Munsell Color Chart NewWindsor NY USA 1990
[20] H O Hartley ldquoThe maximum F-ratio as a short cut test forheterogeneity of variancesrdquo Biometrika vol 37 pp 308ndash3121950
[21] L Pecetti P Annicchiarico and A B Damania ldquoBiodiversity ina germplasm collection of durumwheatrdquo Euphytica vol 60 no3 pp 229ndash238 1992
[22] L Pecetti and A B Damania ldquoGeographic variation intetraploid wheat (Triticum turgidum ssp turgidum convardurum) landraces from two provinces in Ethiopiardquo GeneticResources and Crop Evolution vol 43 no 5 pp 395ndash407 1996
[23] S K Jain C O Qualset G M Bhatt and K K Wu ldquoGeo-graphical patterns of phenotypic diversity in a world collectionof durumwheatsrdquoCrop Science vol 15 no 5 pp 700ndash704 1975
[24] K K Nkongolo L Chinthu M Malusi and Z Vokhiwa ldquoPar-ticipatory variety selection and characterization of sorghum(Sorghum bicolour (L) Moench) elite accessions from Malaw-ian gene pool using farmer and breeder knowledgerdquo AfricanJournal of Agricultural Research vol 3 pp 273ndash283 2008
[25] M B Semira ldquoInnovative methods and metrics for agricul-ture and nutrition actions soil quality and human nutritionin Ethiopiardquo Leverhulme Center for Integrative Research onAgriculture and Health (LCIRAH) 2017
[26] I A Khan A Khatri M A Siddiqui G S Nizamani and SRaza ldquoPerformance of promising sugarcane clone for yield andquality traits in different ecological zones of Sindhirdquo PakistanJournal of Botany vol 36 no 1 pp 83ndash92 2004
[27] M F Perera M E Arias D Costilla et al ldquoGenetic diversityassessment and genotype identification in sugarcane based onDNAmarkers andmorphological traitsrdquo Euphytica vol 185 no3 pp 491ndash510 2012
[28] C G Neil M Kay and C C Jack ldquoDiversity among mainlandUSA sugarcane cultivars examined by SSR genotypingrdquo Journalof the American Society of Sugar Cane Technologists vol 29 pp36ndash52 2009
[29] S SchenckMWCrepeauK KWu P HMooreQ Yu andRMing ldquoGenetic diversity and relationships in native HawaiianSaccharum officinarum sugarcanerdquo Journal of Heredity vol 95no 4 pp 327ndash331 2004
[30] R Gebretsadik H Shimelis M D Laing P Tongoona andN Mandefro ldquoA diagnostic appraisal of the sorghum farmingsystem and breeding priorities in Striga infested agro-ecologiesof Ethiopiardquo Agricultural Systems vol 123 pp 54ndash61 2014
[31] L E V Cesar A T Bruzi J A R Nunes et al ldquoGeneticdivergence of sugarcane varieties collected in the region ofLavras Minas Gerais Brazilrdquo Genetics and Molecular Researchvol 14 no 4 pp 14043ndash14049 2015
[32] C Grenier P J Bramel J A Dahlberg et al ldquoSorghums of theSudan analysis of regional diversity and distributionrdquo GeneticResources and Crop Evolution vol 51 no 5 pp 489ndash500 2004
[33] N V Nair R Balakrishnan and T V Sreenivasan ldquoVariabilityfor quantitative traits in exotic hybrid germplasm of sugarcanerdquoGenetic Resources and Crop Evolution vol 45 no 5 pp 459ndash464 1998
[34] R Nagarajan S Alarmelu and R M Shanthi ldquoStudies onvariation in interspecific hybrids of Saccharumrdquo Sugar Tech vol2 no 3 pp 42ndash46 2000
18 Advances in Agriculture
[35] D N Kamat and J R P Singh ldquoVariability in Sugarcane underrain fed conditionrdquo Sugar Technology Short Communicationsvol 3 no 1-2 pp 65ndash67 2001
[36] B S Rishipal A S M Chaudhary and S P Kadian ldquoStudieson character interrelationship in clonal generation of sugarcane(Saccharum complex hybrids)rdquo Indian Sugar vol 47 no 11 pp907ndash911 1998
[37] S Thippeswamy S T Kajjidoni P M Salimath and J V GoudldquoCorrelation and path analysis for cane yield juice quality andtheir component traits in sugarcanerdquo Short Communication vol5 no 1-2 pp 65ndash72 2003
[38] P S Verma S Pal and N K Karma ldquoGenetic variability andcorrelation studies in sugarcanerdquo Indian Sugar vol 49 pp 125ndash128 1999
[39] H A Esechie J W Maranville and W M Ross ldquoRelationshipof stalk morphology and chemical composition to lodgingresistance in sorghumrdquo Crop Science vol 17 no 4 p 609 1977
[40] P K Das B C Jena and N Nayak ldquoCharacter association andpath analysis of sugar yield in sugarcanerdquo Indian Sugar vol 46no 10 pp 805ndash808 1997
[41] K J Sreekumar M Kuriakose M Thomas D Alexander andS Santhakumari ldquoVariability heritability and correlation onstudies on the yield and quality characters of sugarcanerdquo IndianSugar vol 44 no 4 pp 243ndash249 1994
[42] J R Harlan J M de Wet and E G Price ldquoComparativeevolution of cerealsrdquo Evolution vol 27 no 2 pp 311ndash325 1973
[43] A Elings ldquoDurum wheat landraces from syria II patterns ofvariationrdquo Euphytica vol 54 no 3 pp 231ndash243 1991
[44] K Tato ldquoRainfall in Ethiopiardquo Ethiopian Geographical Journalvol 2 pp 28ndash36 1964
[45] A B L Stemler J R Harlan and J M J deWet ldquoThe sorghumsof ethiopiardquo Economic Botany vol 31 no 4 pp 446ndash460 1977
[46] P Annicchiarico L Pecetti and A B Damania ldquoRelationshipsbetween phenotypic variation and climatic factors at collectingsites in durum wheat landracesrdquo Hereditas vol 122 no 2 pp163ndash167 1995
[47] A B Damania L Pecetti C O Qualset and B O HumeidldquoDiversity and geographic distribution of adaptive traits inTriticum turgidum L (durum group) wheat landraces fromTurkeyrdquoGenetic Resources and Crop Evolution vol 43 no 5 pp409ndash422 1996
[48] A Ayana and E Bekele ldquoGeographical patterns of morpho-logical variation in sorghum (Sorghum bicolor (L) Moench)germplasm from Ethiopia and Eritrea quantitative charactersrdquoEuphytica vol 115 no 2 pp 91ndash104 2000
[49] D S Falconer Introduction to Quantitative Genetics JohnWilley and Sons Inc New York NY USA 3rd edition 1993
[50] R S Thorpe ldquoBiochemical analysis of geographical variationand racial affinitiesrdquo Biology Review vol 51 pp 407ndash452 1976
[51] M Farooq ldquoMorphological and agricultural characteristicsof sugarcane in relation to mechanized agriculture at NARCIslamabadrdquo Portu Rico Journal vol 1 pp 51ndash61 1989
[52] M Akhtar N N Elahi and M Ashraf ldquoEvaluation of exoticsugarcane varieties for agronomic characters and productivityrdquoPakistan Journal of Biological Sciences vol 4 no 1 pp 37ndash402001
[53] F R Piscitelli ldquoPrincipals caracteres exomorfologicos de cul-tures de cana de azucarrdquo TUC Iindustraly Agricos vol 7 pp 49ndash57 1994 (Spanish)
[54] A H D Brown and B S Weir ldquoMeasuring genetic variability inplant populationsrdquo in Isozyme in Plant Genetics Breeding S DTanksley and T J Orton Eds part A Elsevier Science 1983
[55] R Balakrishnan N V Nair and T V Sreenivasan ldquoA methodfor establishing a core collection of Saccharum officinarum Lgermplasm based on quantitative-morphological datardquo GeneticResources and Crop Evolution vol 47 no 1 pp 1ndash9 2000
[56] C A Nestor and B Q Reynaldo Sugarcane Variety Improve-ment in Southeast Asia and the Pacific for Enhanced andSustainable Productivity ndash Germplasm and Disease IndexingComponents Institute of Plant Breeding (IPB) College ofAgriculture (CA) UP Los Banos (UPLB) 2003
[57] A Maqbool J Muhammad and A Sagheer ldquoAgronomic traitsand morphological characteristics of some exotic varieties ofsugarcanerdquoPakistan Journal of Agricultural Research vol 19 no4 2006
This is in agreement with previous studies on sugarcanegermplasm [33ndash35]
This is the first study to report information on geo-graphical pattern of variation in Ethiopian local sugarcanegermplasm which has been lacking so far Our results pro-vided experimental evidence on occurrence of geographicalvariation and significant within-region variation where it washigh in the regions such as Amhara Benshangul-Gumz andSNNPR
The overall patterns of similarity or difference betweenregions seemed to depend on environmental factors such asrainfall temperature length of growing season and altitudeWide variability of millable stalk count at harvest singlecane weight and plant height was observed among regionswhere these characters demonstrated high correlation withcane and sugar yield (Table 7) Therefore breeders can utilizeaccessions of regions showing variability for these charactersin selection programs and to design breeding strategies toproduce varieties with best commercial merits
Bivariate Statistics Correlation coefficients worked out onthe 21 quantitative characters are shown for the entire data(Table 7) between regions (Table 8) and within regions(data not shown) Results of the association studies for theentire data showed that cane and sugar yield had highlysignificant positive correlation with most of the quantitativestalk characters and juice quality parameters However theassociation of tiller counts 4 and 5 months after plantingstalk count 10 months after planting millable stalk countat harvest single cane weight and plant height with caneand sugar yield was stronger suggesting these componentcharacters asmain contributing factor to cane and sugar yieldIn fact sugar yield is the product of cane yield and sucrose per-cent cane surely being influenced with components affectingcane yield Similar trend was reported by earlier sugarcaneworkers [36 37] In sugarcane the cane and sugar yields areconsidered to be the complex characters The information onthe phenotypic interrelationship of cane yield and sugar yieldwith their component character per se would be of immensehelp to sugarcane breeders
Stem diameter had significant positive association withboth cane and sugar yield indicating the significance ofthis trait in improving both cane and sugar yields Similarreports have beenmade by earlier sugarcane workers [37 38]Stalk diameter was also positively and significantly correlatedwith plant height It is one of the traits that are related tolodging resistance [39] and the positive associations withheight would help in reducing the chance of lodging as heightincreases Among the physiological attributes included in thestudy leaf area had highly significant positive associationwith cane yield Similar observation was also made in [37]Sucrose percent which is an important juice quality attributehad highly significant positive association with sugar yieldand simultaneously cane yield also had highly significantassociation with sugar yield Based on the magnitude ofcorrelation coefficient values it can be said that cane yield ismuch more important than sucrose percent in determiningthe sugar yield and the same has also been reported by othersugarcane workers [37 40] Similar to observations made in
[37 41] results of the current study showed that all the qualityparameters like sucrose percent pol percent brix percentand purity percent were significantly correlated in positivedirection This indicated any of these juice quality traitscould be considered for selection leading to the simultaneousimprovement in the remaining quality traits
Ten of the 21 characters also showed significant positivecorrelations with altitude of the collection sites (Table 7) Asdescribed in [42] ecological characteristics have influencedthe genotypic constitution of landraces during domesticationand hence a relationship exists between the agroecology inthe collection site and the morphological characteristics ofthe landraces Thus positive correlation between collectionsite variables and plant characteristics would imply that thevariation between accessionsmay be related to agroecologicalvariations among the collection sites [43]
The correlation coefficient between number of internodesand altitude was negative and significant indicating thatother environmental factors (other than altitude) andornonenvironmental factorsmight account for the variation forthis particular character Since temperature decreases withan increase of altitude in Ethiopia [44 45] it is more likelythat temperature has exerted strong selection pressure innumber of internodes in the genotypes Information on therelationships between environmental factors of the collect-ing sites and morphophysiological variation of germplasmcould enhance the understanding of evolutionary adaptivepatterns which could assist germplasm collectors and users[46]
Character associations between region and within regionalso followed similar fashion like that of the entire dataNumber of sprout tiller and stalk counts and sugar qualityparameters showed significant positive correlations with caneand sugar yield
Though not significant and strong the correlation of leaflength and width and leaf area (leaf area = leaf length Xleaf width X 075) with cane and sugar yield was negativein the interregion correlations This kind of relationship wasalso exhibited between these leaf characteristics and yieldcomponents like tiller and stalks counts plant height andsingle cane weight and stalk diameter and sugar qualityparameters This may have been attributed to the variationof temperature with altitude and other environmental factorsacross the regions resulting in such type of relationshipSuch relationship also calls for the need to have enough dataon environmental variables of the collection sites such astemperature and rainfall if germplasm collection is requiredtomeet its objectives effectively and efficiently Unfortunatelydata on such useful environmental variables like temperatureand rainfall are lacking in the passport data of the currentEthiopian sugarcane germplasm collections The significanceof environmental factors of the collection sites as veryimportant determinants in structuring morphophysiolog-ical variations was reported in tetraploid wheat [22 4748]
Association of characters among yield its componentsand other economic traits is important for the interpretationof the patterns of variation and making selection in breed-ing program and combining several desirable attributes It
suggests the advantage of a scheme of selection for morethan one character at a time The correlation between char-acters may arise from linkage or from developmental geneticinteractions with orwithout a purely phenotypic componentIt could arise also due to differential phenotypic plasticityof characters themselves Within the limit of experimentalerror and environmental effects high correlation coefficientsbetween characters may show that the characters sharesome common element of genetic control (ie pleiotropylinkage) between genes or else from independently controlledcharacters responding similarly to geographic variation inselection pressures [49] The interregion correlation coef-ficient between the characters measures the consistency oftheir patterns of regional variation while the intraregioncorrelation coefficient measures the association arising fromgenetic factors but is not affected by regional variation[50] Since this study showed significant positive correlationsintraregionally for some character combinations it wouldseem that common genetic control might be playing a role inbringing about correlations between the various charactersIt appeared that different response to regional variationwas playing a greater role than different genetic controlas shown by the many more significant and moderate tohigh correlation coefficients intraregionally than interregion-ally
322 Qualitative Characters The frequency distribution forthe 16 qualitative characters of germplasm samples by regionsin Ethiopia and of the exotic accessions is shown in Supple-mental Table 5 For the purpose of avoiding redundancy andimproving readability the discrete characters are presented bygrouping related traits together
BudCushion andRelativeDegree of BudExtension Evaluationof the presence and absence of bud cushion revealed thatgreat proportion (70) of sugarcane landraces from Ethiopiado not have bud cushion Majority of the sample germplasmin all regions belong to this phenotypic class Approximatelyone-half (53) of the collection have their bud extendedabove the growth ring followed by those (41) having budtouching the growth ring
Relative Bud Shape Among the phenotypic classes for thischaracter tall deltoid was dominant (31) followed by ovate(17) and narrow ovate (12) While local accessions inmost regions had tall deltoid bud shape 50 of the intro-duced accessions belong to two phenotypic classes roundwith central germpore (30) and 20 having pentagonalbud shape Equal proportions (19) of the samples fromBenshangul-Gumz were Ovate and Ovate with emarginatebasal wing
Canopy Structure Relative Plant Erectness Colour of theLeaves and Colour of the Exposed Rind Out of the eightphenotypic classes observed compact tip droopy was themost frequent (35) Open semidroopy and open droopyclasses had fairly equal distribution With regard to relativeplant erectness seventy-five percent of the germplasm werefound to fall in twophenotypic classes erect (41) and almost
erect (34) for the whole region and country Most of theaccessions (90) showed light green or green leaf colourbut 20 from introduced collections were greenish yellowAmong phenotypic classes for colour of the exposed rindbrownish yellow yellow brown and yellowish green were themost frequent with fairly equal distribution
Relative Shape of Dewlap Double crescent had larger fre-quency (22) while squarish descending and flaring typeof dewlap shape had fairly equal proportions Seventeenpercent of germplasm from Tigray region had squarishdeltoid dewlap shape
Relative Degree of Internode Alignment and Internode ShapeForty-four percent of the sample germplasm were slightlyzigzag while there was almost equal frequency for straightand zigzag internode alignment for the entire region andcountry Straight or nearly straight (slightly zigzag) internodealignment of cane stalks is a very important character formechanized farming and postharvest handling in sugarcane[51] Forty percent of the collections were observed havingconcave-convex type of internode shape while 29 wascylindrical and 14 conoidal Other classes had fairly equalproportion
Type of Auricle and Relative Shape of Ligule An equalproportion of the germplasm totally constituting 84 ofthe collection belong to four phenotypic classes namelythose with no auricle and others with transitional and shortand lanceolate auricles Sixteen percent of the collectionsalso showed equal frequency for the remaining classesBenshangul-Gumz and Tigray regions had each equal 13accessions with falcate type of auricle Sixty-seven percentof the collections belong to three ligule shape classes cres-cent with lozenge (27) broad-crescent (21) and linear-crescent (19) The germplasm had almost equal proportionfor the other phenotypic classes of this character Nineteenpercent of the accessions fromBenshangul-Gumz region haddeltoid shape of ligule
Stalk Corky Cracks Corky Patches Growth Cracks and BudGroove Eighty-nine percent of the germplasm across regionshad no stalk corky cracks with only eleven percent exhibitingthis character All the germplasm in Gambella Harari andSomali had no stalk corky cracks On the other hand eighty-one percent of the germplasm for the whole region showedthe presence of stalk corky patches All the local germplasmin Benshangul-Gumz Gambella and Tigray had stalk corkypatches whereas 75 of the germplasm from Harari Region67 from Somali Region and 24 from Oromia Regionhad no stalk corky patches In the entire region 75 ofthe germplasm had no stalk growth cracks with only 25showing this character Sixty-three percent of the germplasmwere found to have bud groove Howevermost of the samplesin Harari (75) Amhara (63) and Oromia (53) Regionshad no bud groove
The results of this study indicated the wide distributionof phenotypic classes for the characters considered whichindicates the existence of different races and combination
Advances in Agriculture 15
of races in local genotypes of different regions in EthiopiaThis diversity of germplasmbased onphenotypicmarkers canhelp sugarcane breeders in identifying the identity of geno-types as well as maintaining genetic diversity Differencesin morphological characters of different sugarcane varietieshave been reported [52] Piscitelli [53] showed importantexomorphological qualitative characters of sugarcane varietythat are not influenced by environmental factors and thus canbe used as a selection tool in any breeding program
The amount of phenotypic diversity estimates based onthe ShannonndashWeaver diversity index (H1015840) are shown inTable 9 The 16 characters differed in their distribution aswell as the amount of variation Individual traits showed adifferent pattern of variation among accessions Estimatesof H1015840 for individual characters varied from 049 for stalkcorky cracks to 100 for bud groove with an overall meanof 080 Most of the characters were highly polymorphicwhereas relative plant erectness and presence or absenceof stalk corky patches scored moderate values The highdiversity values for the characters showed a wide variabilityamong genotypes According to [54] this index is also usedin genetic resource studies as a convenient measure of bothallelic richness and allelic evenness when using geneticdata
The highest diversity computed was on presence orabsence of bud groove with a diversity index of 100indicating that this character is very variable in the localsugarcane germplasm On the other hand the descriptorcomputed with the lowest diversity index value (049) wasthe presence or absence of stalk corky cracks indicatingthat this character is selected in the development of thegenotypes In agreement with the present study [55] workingon S officinarum accessions from the world collection ofsugarcane germplasm obtained a low index of 025 and 035for presence or absence of stalk corky patches and cracksrespectively
Their report also showed high diversity index values forinternode alignment growth cracks internode shape budcushion bud extension bud groove and ligule shape Sim-ilarly [56] experimenting on field collections of sugarcaneaccessions in Philippine reported high ShannonndashWeaverdiversity for stalk growth cracks bud cushion stalk corkypatch stalk corky cracks bud extension and auricle outershape Unlike the present study they have reported high indexfor stalk corky cracksMedium index values were reported forstalk internode shape and low index values for ligule shapeand bud shape High variability of sugarcane varieties fordewlap shape was also reported in [57]
The H1015840 pooled across characters by region ranged from079 in Oromiya to 085 in Tigray with an overall aver-age of 081 (Table 8) Most of the regions showed highShannonndashWeaver diversity index The populations that hadthe highest H1015840 were that of Tigray and introduction The H1015840averaged over regions for different characters was found torange from 044 for presence or absence of stalk corky cracksto 094 for relative degree of internode alignment and budgroove with an overall average of 081 (Table 9)
Assessment of the ShannonndashWeaver index also showedsensitivity to both the number of descriptor classes and the
distribution within phenotypic classes in each region Amongregions the mean values for the ShannonndashWeaver index arenot significantly different but differences were found whenindividual characters were considered These observationssuggest that some regions have high diversity for particulartraits while in other regions selective pressures might havereduced the variation to certain genotypes
4 Conclusions
The large number of landraces observed during the cur-rent study demonstrates the existence of diverse geneticresources of sugarcane in Ethiopia The landraces wouldserve to broaden the genetic base of the available sugarcanegermplasm Generally distribution of the overall phenotypicdiversity among sugarcane landraces in Ethiopia is unevenOur results showed that there was a wide range of variationthat existed in the local sugarcane landraces studied both atregional and within region levels Selection and differenti-ation of types might have taken place along a geographicalpattern perhaps associated with climate and use The presentstudy contributes to updating sugarcane descriptors adoptedfrom USDA-ARS as well as Bioversity passport data forthe future collection and evaluation As Ethiopian SugarCorporation is in the course of launching breeding programthe observed substantial variation of landraces would enablesugarcane breeders to design and practice breeding andselection programs to improve the crop
Data Availability
The data used to support the findings of this study areavailable from the corresponding author upon request
Conflicts of Interest
The authors declare that they have no conflicts of interest
Acknowledgments
The authors are grateful for the financial grant of SugarCorporation of Ethiopia Many thanks are due to EthiopianBiodiversity Institute (EBI) for its technical advice duringcollection of the local sugarcane genotypes Sincere thanksare forwarded to Ethiopian smallholder farmers who pre-served sugarcane germplasm and shared their knowledgeand experience Regional Zonal and District (ldquoWeredardquo)Agriculture Bureaus across Ethiopia are acknowledged fortheir support during the study
Supplementary Materials
These supplementary materials contain different tables sup-porting the results of the study These include passport dataof sugarcane germplasm collected and related informationon the materials local sugarcane germplasm collected andused formorphological diversity study introduced sugarcane
16 Advances in Agriculture
Table9Sh
anno
n-WeaverD
iversityIndices(H1015840)a
ndmeanandsta
ndarderror(plusmnS
E)of
land
racesineach
region
basedon
16discretecharacters
Characters
Amha
raBe
nsha
ngul-G
umz
Introd
uced
Oromia
SNNP
Tigray
ELb
Total
Presence
orabsenceo
fbud
cushion(BUDCUSH
ION)
083
095
095
089
082
098
090
088
Relativ
edegreeo
fbud
extension(BUDEX
TEND)
077
079
093
074
072
087
080
078
Relativ
ebud
Shape(BU
DSH
APE
)078
096
089
081
085
078
084
078
Canop
ystructure(C
ANOPY
)089
098
095
081
078
096
089
084
Relativ
eshape
ofdewlap(D
EWLA
PSHAP)
087
094
092
090
082
096
090
086
Relativ
eplant
erectness(ER
ECT)
080
082
050
067
071
088
073
064
Relativ
edegreeo
finterno
dealignm
ent(IN
ALIGN)
096
076
098
097
098
100
094
098
Relativ
einterno
deshape(IN
SHAPE
)086
069
095
077
083
091
083
082
Color
oftheleaves(LE
AFC
OLO
R)087
071
090
064
074
071
076
076
Type
ofou
tera
uricle(A
URI
CLE
OUT)
080
092
087
079
083
093
086
082
Relativ
eshape
ofligule(LIGSH
APE
)086
094
082
085
090
083
087
083
Presence
orabsenceo
fstalkcorkyc
racks(ST
ALK
CORK
C)
044
054
035
055
051
025
044
049
Presence
orabsenceo
fstalkcorkyp
atches(STA
LKCORK
P)066
000
072
079
059
087
060
069
Presence
orabsenceo
fstalkgrow
thcracks
(STA
LKCRA
CK)
066
100
088
062
088
099
084
080
Color
ofthee
xposed
rind(R
INDCOLE
)085
096
089
084
086
076
086
085
Presence
orabsenceo
fbud
groo
ve(BUDGRO
OVE)
095
100
072
100
100
099
094
100
Means
080plusmn0
03
081plusmn0
06
083plusmn0
04
079plusmn0
03
080plusmn0
03
085plusmn0
05
081plusmn0
03
080
b EL=
averageo
fentire
locatio
n
Advances in Agriculture 17
varieties used for morphological diversity study and fre-quency distribution for 16 qualitative characters in sugarcaneby regions in Ethiopia and country of sources of introducedgenotypes (Supplementary Materials)
References
[1] ldquoSpreading the word about clean solutions from sugarcanrdquoAccessed 04072018 httpwwwsugarcaneorg
[2] Central Statistics Agency (CSA) ldquoAgricultural sample surveyArea and production of major crops private peasant holdingsrdquoStatistical Bulletin Addis Ababa Ethiopia 2017
[3] T Esayas Exploration and Collection Characterization GeneticDiversity Analysis and Association of Traits for Yield and YieldComponents of Sugarcane (Saccharumspp) in Ethiopia [MasterThesis] Haramaya University Dire Dawa Ethiopia 2014
[4] J R Harlan Crops and Man American Society of AgronomyCrop Science Society of America Madison WI USA 2ndedition 1992
[5] R W Allard Principles of Plant Breeding JohnWilley and SonsNew York NY USA 2nd edition 1999
[6] B T Roach and J Daniels ldquoA review of the origin and improve-ment of sugarcanerdquo in Proceedings of Coper sugar InternationalSugarcane Workshop pp 1ndash32 1987
[7] P P Patil S S Patil and D U Patel ldquoGenetic diversity in sug-arcane (Saccharum spp Complex)rdquo Journal of Pharmacognosyand Phytochemistry vol 6 no 6 pp 570ndash573 2017
[8] R B Singh B Singh andR K Singh ldquoStudy of genetic diversityof sugarcane (Saccharum) species and commercial varietiesthrough TRAP molecular markersrdquo Indian Journal of PlantPhysiology vol 22 no 3 pp 332ndash338 2017
[9] R W Allard ldquoPopulation structure and sampling methodsrdquo inGenetic Resources in PlantsmdashTheir Exploration and Conserva-tion O H Frankel and E Bennett Eds IBP Handbook no 11pp 97ndash107 FA Davis Company Philadelphia PA USA 1970
[10] D R Marshall and A H D Brown ldquoOptimum samplingstrategies in genetic conservationrdquo inCrop Genetic Resources forToday and Tomorrow O H Frankel and J G Hawkes Eds pp53ndash80 Cambridge University Press 1975
[11] J G Hawkes Crop Genetic Resources Field Collection ManualPudoc Wagnningen Netherland 1980
[12] J G Hawkes The Diversity of Crop Plants Harvard UniversityPress London England 1983
[13] B P K Yerima Manual for Good Laboratory Practice Foodand Agriculture Organization of United Nation Addis AbabaEthiopia 1993
[14] A Walkley and I A Black ldquoAn examination of the degtjareffmethod for determining soil organic matter and a proposedmodification of the chromic acid titrationmethodrdquo Soil Sciencevol 37 no 1 pp 29ndash38 1934
[15] S R Olsen and L E Sommers ldquoPhosphorusrdquo in AgronomyMonograph A L Page et al Ed Methods of soil analysis part2 pp 403ndash430 ASA and SSSAMadisonWI USA 2nd edition1982
[16] R H Bray and L T Kurtz ldquoDetermination of total organic andavailable forms of phosphorus in soilsrdquo Soil Science vol 59 no1 pp 39ndash46 1945
[17] S Sahlemedhin andB Taye ldquoProcedures for soil and plant anal-ysisrdquo Tech Rep 74 National Soil Research Center EthiopianAgricultural Research Organization 2000
[18] F C Stickler S Weaden and AW Pauli ldquoLeaf area determina-tion in grain sorghumrdquo Journal of Agronomy and Crop Sciencevol 53 pp 187-188 1961
[19] Munsell Color ChartThe Munsell Book of Color Matte Collec-tion Munsell Color Chart NewWindsor NY USA 1990
[20] H O Hartley ldquoThe maximum F-ratio as a short cut test forheterogeneity of variancesrdquo Biometrika vol 37 pp 308ndash3121950
[21] L Pecetti P Annicchiarico and A B Damania ldquoBiodiversity ina germplasm collection of durumwheatrdquo Euphytica vol 60 no3 pp 229ndash238 1992
[22] L Pecetti and A B Damania ldquoGeographic variation intetraploid wheat (Triticum turgidum ssp turgidum convardurum) landraces from two provinces in Ethiopiardquo GeneticResources and Crop Evolution vol 43 no 5 pp 395ndash407 1996
[23] S K Jain C O Qualset G M Bhatt and K K Wu ldquoGeo-graphical patterns of phenotypic diversity in a world collectionof durumwheatsrdquoCrop Science vol 15 no 5 pp 700ndash704 1975
[24] K K Nkongolo L Chinthu M Malusi and Z Vokhiwa ldquoPar-ticipatory variety selection and characterization of sorghum(Sorghum bicolour (L) Moench) elite accessions from Malaw-ian gene pool using farmer and breeder knowledgerdquo AfricanJournal of Agricultural Research vol 3 pp 273ndash283 2008
[25] M B Semira ldquoInnovative methods and metrics for agricul-ture and nutrition actions soil quality and human nutritionin Ethiopiardquo Leverhulme Center for Integrative Research onAgriculture and Health (LCIRAH) 2017
[26] I A Khan A Khatri M A Siddiqui G S Nizamani and SRaza ldquoPerformance of promising sugarcane clone for yield andquality traits in different ecological zones of Sindhirdquo PakistanJournal of Botany vol 36 no 1 pp 83ndash92 2004
[27] M F Perera M E Arias D Costilla et al ldquoGenetic diversityassessment and genotype identification in sugarcane based onDNAmarkers andmorphological traitsrdquo Euphytica vol 185 no3 pp 491ndash510 2012
[28] C G Neil M Kay and C C Jack ldquoDiversity among mainlandUSA sugarcane cultivars examined by SSR genotypingrdquo Journalof the American Society of Sugar Cane Technologists vol 29 pp36ndash52 2009
[29] S SchenckMWCrepeauK KWu P HMooreQ Yu andRMing ldquoGenetic diversity and relationships in native HawaiianSaccharum officinarum sugarcanerdquo Journal of Heredity vol 95no 4 pp 327ndash331 2004
[30] R Gebretsadik H Shimelis M D Laing P Tongoona andN Mandefro ldquoA diagnostic appraisal of the sorghum farmingsystem and breeding priorities in Striga infested agro-ecologiesof Ethiopiardquo Agricultural Systems vol 123 pp 54ndash61 2014
[31] L E V Cesar A T Bruzi J A R Nunes et al ldquoGeneticdivergence of sugarcane varieties collected in the region ofLavras Minas Gerais Brazilrdquo Genetics and Molecular Researchvol 14 no 4 pp 14043ndash14049 2015
[32] C Grenier P J Bramel J A Dahlberg et al ldquoSorghums of theSudan analysis of regional diversity and distributionrdquo GeneticResources and Crop Evolution vol 51 no 5 pp 489ndash500 2004
[33] N V Nair R Balakrishnan and T V Sreenivasan ldquoVariabilityfor quantitative traits in exotic hybrid germplasm of sugarcanerdquoGenetic Resources and Crop Evolution vol 45 no 5 pp 459ndash464 1998
[34] R Nagarajan S Alarmelu and R M Shanthi ldquoStudies onvariation in interspecific hybrids of Saccharumrdquo Sugar Tech vol2 no 3 pp 42ndash46 2000
18 Advances in Agriculture
[35] D N Kamat and J R P Singh ldquoVariability in Sugarcane underrain fed conditionrdquo Sugar Technology Short Communicationsvol 3 no 1-2 pp 65ndash67 2001
[36] B S Rishipal A S M Chaudhary and S P Kadian ldquoStudieson character interrelationship in clonal generation of sugarcane(Saccharum complex hybrids)rdquo Indian Sugar vol 47 no 11 pp907ndash911 1998
[37] S Thippeswamy S T Kajjidoni P M Salimath and J V GoudldquoCorrelation and path analysis for cane yield juice quality andtheir component traits in sugarcanerdquo Short Communication vol5 no 1-2 pp 65ndash72 2003
[38] P S Verma S Pal and N K Karma ldquoGenetic variability andcorrelation studies in sugarcanerdquo Indian Sugar vol 49 pp 125ndash128 1999
[39] H A Esechie J W Maranville and W M Ross ldquoRelationshipof stalk morphology and chemical composition to lodgingresistance in sorghumrdquo Crop Science vol 17 no 4 p 609 1977
[40] P K Das B C Jena and N Nayak ldquoCharacter association andpath analysis of sugar yield in sugarcanerdquo Indian Sugar vol 46no 10 pp 805ndash808 1997
[41] K J Sreekumar M Kuriakose M Thomas D Alexander andS Santhakumari ldquoVariability heritability and correlation onstudies on the yield and quality characters of sugarcanerdquo IndianSugar vol 44 no 4 pp 243ndash249 1994
[42] J R Harlan J M de Wet and E G Price ldquoComparativeevolution of cerealsrdquo Evolution vol 27 no 2 pp 311ndash325 1973
[43] A Elings ldquoDurum wheat landraces from syria II patterns ofvariationrdquo Euphytica vol 54 no 3 pp 231ndash243 1991
[44] K Tato ldquoRainfall in Ethiopiardquo Ethiopian Geographical Journalvol 2 pp 28ndash36 1964
[45] A B L Stemler J R Harlan and J M J deWet ldquoThe sorghumsof ethiopiardquo Economic Botany vol 31 no 4 pp 446ndash460 1977
[46] P Annicchiarico L Pecetti and A B Damania ldquoRelationshipsbetween phenotypic variation and climatic factors at collectingsites in durum wheat landracesrdquo Hereditas vol 122 no 2 pp163ndash167 1995
[47] A B Damania L Pecetti C O Qualset and B O HumeidldquoDiversity and geographic distribution of adaptive traits inTriticum turgidum L (durum group) wheat landraces fromTurkeyrdquoGenetic Resources and Crop Evolution vol 43 no 5 pp409ndash422 1996
[48] A Ayana and E Bekele ldquoGeographical patterns of morpho-logical variation in sorghum (Sorghum bicolor (L) Moench)germplasm from Ethiopia and Eritrea quantitative charactersrdquoEuphytica vol 115 no 2 pp 91ndash104 2000
[49] D S Falconer Introduction to Quantitative Genetics JohnWilley and Sons Inc New York NY USA 3rd edition 1993
[50] R S Thorpe ldquoBiochemical analysis of geographical variationand racial affinitiesrdquo Biology Review vol 51 pp 407ndash452 1976
[51] M Farooq ldquoMorphological and agricultural characteristicsof sugarcane in relation to mechanized agriculture at NARCIslamabadrdquo Portu Rico Journal vol 1 pp 51ndash61 1989
[52] M Akhtar N N Elahi and M Ashraf ldquoEvaluation of exoticsugarcane varieties for agronomic characters and productivityrdquoPakistan Journal of Biological Sciences vol 4 no 1 pp 37ndash402001
[53] F R Piscitelli ldquoPrincipals caracteres exomorfologicos de cul-tures de cana de azucarrdquo TUC Iindustraly Agricos vol 7 pp 49ndash57 1994 (Spanish)
[54] A H D Brown and B S Weir ldquoMeasuring genetic variability inplant populationsrdquo in Isozyme in Plant Genetics Breeding S DTanksley and T J Orton Eds part A Elsevier Science 1983
[55] R Balakrishnan N V Nair and T V Sreenivasan ldquoA methodfor establishing a core collection of Saccharum officinarum Lgermplasm based on quantitative-morphological datardquo GeneticResources and Crop Evolution vol 47 no 1 pp 1ndash9 2000
[56] C A Nestor and B Q Reynaldo Sugarcane Variety Improve-ment in Southeast Asia and the Pacific for Enhanced andSustainable Productivity ndash Germplasm and Disease IndexingComponents Institute of Plant Breeding (IPB) College ofAgriculture (CA) UP Los Banos (UPLB) 2003
[57] A Maqbool J Muhammad and A Sagheer ldquoAgronomic traitsand morphological characteristics of some exotic varieties ofsugarcanerdquoPakistan Journal of Agricultural Research vol 19 no4 2006
This is in agreement with previous studies on sugarcanegermplasm [33ndash35]
This is the first study to report information on geo-graphical pattern of variation in Ethiopian local sugarcanegermplasm which has been lacking so far Our results pro-vided experimental evidence on occurrence of geographicalvariation and significant within-region variation where it washigh in the regions such as Amhara Benshangul-Gumz andSNNPR
The overall patterns of similarity or difference betweenregions seemed to depend on environmental factors such asrainfall temperature length of growing season and altitudeWide variability of millable stalk count at harvest singlecane weight and plant height was observed among regionswhere these characters demonstrated high correlation withcane and sugar yield (Table 7) Therefore breeders can utilizeaccessions of regions showing variability for these charactersin selection programs and to design breeding strategies toproduce varieties with best commercial merits
Bivariate Statistics Correlation coefficients worked out onthe 21 quantitative characters are shown for the entire data(Table 7) between regions (Table 8) and within regions(data not shown) Results of the association studies for theentire data showed that cane and sugar yield had highlysignificant positive correlation with most of the quantitativestalk characters and juice quality parameters However theassociation of tiller counts 4 and 5 months after plantingstalk count 10 months after planting millable stalk countat harvest single cane weight and plant height with caneand sugar yield was stronger suggesting these componentcharacters asmain contributing factor to cane and sugar yieldIn fact sugar yield is the product of cane yield and sucrose per-cent cane surely being influenced with components affectingcane yield Similar trend was reported by earlier sugarcaneworkers [36 37] In sugarcane the cane and sugar yields areconsidered to be the complex characters The information onthe phenotypic interrelationship of cane yield and sugar yieldwith their component character per se would be of immensehelp to sugarcane breeders
Stem diameter had significant positive association withboth cane and sugar yield indicating the significance ofthis trait in improving both cane and sugar yields Similarreports have beenmade by earlier sugarcane workers [37 38]Stalk diameter was also positively and significantly correlatedwith plant height It is one of the traits that are related tolodging resistance [39] and the positive associations withheight would help in reducing the chance of lodging as heightincreases Among the physiological attributes included in thestudy leaf area had highly significant positive associationwith cane yield Similar observation was also made in [37]Sucrose percent which is an important juice quality attributehad highly significant positive association with sugar yieldand simultaneously cane yield also had highly significantassociation with sugar yield Based on the magnitude ofcorrelation coefficient values it can be said that cane yield ismuch more important than sucrose percent in determiningthe sugar yield and the same has also been reported by othersugarcane workers [37 40] Similar to observations made in
[37 41] results of the current study showed that all the qualityparameters like sucrose percent pol percent brix percentand purity percent were significantly correlated in positivedirection This indicated any of these juice quality traitscould be considered for selection leading to the simultaneousimprovement in the remaining quality traits
Ten of the 21 characters also showed significant positivecorrelations with altitude of the collection sites (Table 7) Asdescribed in [42] ecological characteristics have influencedthe genotypic constitution of landraces during domesticationand hence a relationship exists between the agroecology inthe collection site and the morphological characteristics ofthe landraces Thus positive correlation between collectionsite variables and plant characteristics would imply that thevariation between accessionsmay be related to agroecologicalvariations among the collection sites [43]
The correlation coefficient between number of internodesand altitude was negative and significant indicating thatother environmental factors (other than altitude) andornonenvironmental factorsmight account for the variation forthis particular character Since temperature decreases withan increase of altitude in Ethiopia [44 45] it is more likelythat temperature has exerted strong selection pressure innumber of internodes in the genotypes Information on therelationships between environmental factors of the collect-ing sites and morphophysiological variation of germplasmcould enhance the understanding of evolutionary adaptivepatterns which could assist germplasm collectors and users[46]
Character associations between region and within regionalso followed similar fashion like that of the entire dataNumber of sprout tiller and stalk counts and sugar qualityparameters showed significant positive correlations with caneand sugar yield
Though not significant and strong the correlation of leaflength and width and leaf area (leaf area = leaf length Xleaf width X 075) with cane and sugar yield was negativein the interregion correlations This kind of relationship wasalso exhibited between these leaf characteristics and yieldcomponents like tiller and stalks counts plant height andsingle cane weight and stalk diameter and sugar qualityparameters This may have been attributed to the variationof temperature with altitude and other environmental factorsacross the regions resulting in such type of relationshipSuch relationship also calls for the need to have enough dataon environmental variables of the collection sites such astemperature and rainfall if germplasm collection is requiredtomeet its objectives effectively and efficiently Unfortunatelydata on such useful environmental variables like temperatureand rainfall are lacking in the passport data of the currentEthiopian sugarcane germplasm collections The significanceof environmental factors of the collection sites as veryimportant determinants in structuring morphophysiolog-ical variations was reported in tetraploid wheat [22 4748]
Association of characters among yield its componentsand other economic traits is important for the interpretationof the patterns of variation and making selection in breed-ing program and combining several desirable attributes It
suggests the advantage of a scheme of selection for morethan one character at a time The correlation between char-acters may arise from linkage or from developmental geneticinteractions with orwithout a purely phenotypic componentIt could arise also due to differential phenotypic plasticityof characters themselves Within the limit of experimentalerror and environmental effects high correlation coefficientsbetween characters may show that the characters sharesome common element of genetic control (ie pleiotropylinkage) between genes or else from independently controlledcharacters responding similarly to geographic variation inselection pressures [49] The interregion correlation coef-ficient between the characters measures the consistency oftheir patterns of regional variation while the intraregioncorrelation coefficient measures the association arising fromgenetic factors but is not affected by regional variation[50] Since this study showed significant positive correlationsintraregionally for some character combinations it wouldseem that common genetic control might be playing a role inbringing about correlations between the various charactersIt appeared that different response to regional variationwas playing a greater role than different genetic controlas shown by the many more significant and moderate tohigh correlation coefficients intraregionally than interregion-ally
322 Qualitative Characters The frequency distribution forthe 16 qualitative characters of germplasm samples by regionsin Ethiopia and of the exotic accessions is shown in Supple-mental Table 5 For the purpose of avoiding redundancy andimproving readability the discrete characters are presented bygrouping related traits together
BudCushion andRelativeDegree of BudExtension Evaluationof the presence and absence of bud cushion revealed thatgreat proportion (70) of sugarcane landraces from Ethiopiado not have bud cushion Majority of the sample germplasmin all regions belong to this phenotypic class Approximatelyone-half (53) of the collection have their bud extendedabove the growth ring followed by those (41) having budtouching the growth ring
Relative Bud Shape Among the phenotypic classes for thischaracter tall deltoid was dominant (31) followed by ovate(17) and narrow ovate (12) While local accessions inmost regions had tall deltoid bud shape 50 of the intro-duced accessions belong to two phenotypic classes roundwith central germpore (30) and 20 having pentagonalbud shape Equal proportions (19) of the samples fromBenshangul-Gumz were Ovate and Ovate with emarginatebasal wing
Canopy Structure Relative Plant Erectness Colour of theLeaves and Colour of the Exposed Rind Out of the eightphenotypic classes observed compact tip droopy was themost frequent (35) Open semidroopy and open droopyclasses had fairly equal distribution With regard to relativeplant erectness seventy-five percent of the germplasm werefound to fall in twophenotypic classes erect (41) and almost
erect (34) for the whole region and country Most of theaccessions (90) showed light green or green leaf colourbut 20 from introduced collections were greenish yellowAmong phenotypic classes for colour of the exposed rindbrownish yellow yellow brown and yellowish green were themost frequent with fairly equal distribution
Relative Shape of Dewlap Double crescent had larger fre-quency (22) while squarish descending and flaring typeof dewlap shape had fairly equal proportions Seventeenpercent of germplasm from Tigray region had squarishdeltoid dewlap shape
Relative Degree of Internode Alignment and Internode ShapeForty-four percent of the sample germplasm were slightlyzigzag while there was almost equal frequency for straightand zigzag internode alignment for the entire region andcountry Straight or nearly straight (slightly zigzag) internodealignment of cane stalks is a very important character formechanized farming and postharvest handling in sugarcane[51] Forty percent of the collections were observed havingconcave-convex type of internode shape while 29 wascylindrical and 14 conoidal Other classes had fairly equalproportion
Type of Auricle and Relative Shape of Ligule An equalproportion of the germplasm totally constituting 84 ofthe collection belong to four phenotypic classes namelythose with no auricle and others with transitional and shortand lanceolate auricles Sixteen percent of the collectionsalso showed equal frequency for the remaining classesBenshangul-Gumz and Tigray regions had each equal 13accessions with falcate type of auricle Sixty-seven percentof the collections belong to three ligule shape classes cres-cent with lozenge (27) broad-crescent (21) and linear-crescent (19) The germplasm had almost equal proportionfor the other phenotypic classes of this character Nineteenpercent of the accessions fromBenshangul-Gumz region haddeltoid shape of ligule
Stalk Corky Cracks Corky Patches Growth Cracks and BudGroove Eighty-nine percent of the germplasm across regionshad no stalk corky cracks with only eleven percent exhibitingthis character All the germplasm in Gambella Harari andSomali had no stalk corky cracks On the other hand eighty-one percent of the germplasm for the whole region showedthe presence of stalk corky patches All the local germplasmin Benshangul-Gumz Gambella and Tigray had stalk corkypatches whereas 75 of the germplasm from Harari Region67 from Somali Region and 24 from Oromia Regionhad no stalk corky patches In the entire region 75 ofthe germplasm had no stalk growth cracks with only 25showing this character Sixty-three percent of the germplasmwere found to have bud groove Howevermost of the samplesin Harari (75) Amhara (63) and Oromia (53) Regionshad no bud groove
The results of this study indicated the wide distributionof phenotypic classes for the characters considered whichindicates the existence of different races and combination
Advances in Agriculture 15
of races in local genotypes of different regions in EthiopiaThis diversity of germplasmbased onphenotypicmarkers canhelp sugarcane breeders in identifying the identity of geno-types as well as maintaining genetic diversity Differencesin morphological characters of different sugarcane varietieshave been reported [52] Piscitelli [53] showed importantexomorphological qualitative characters of sugarcane varietythat are not influenced by environmental factors and thus canbe used as a selection tool in any breeding program
The amount of phenotypic diversity estimates based onthe ShannonndashWeaver diversity index (H1015840) are shown inTable 9 The 16 characters differed in their distribution aswell as the amount of variation Individual traits showed adifferent pattern of variation among accessions Estimatesof H1015840 for individual characters varied from 049 for stalkcorky cracks to 100 for bud groove with an overall meanof 080 Most of the characters were highly polymorphicwhereas relative plant erectness and presence or absenceof stalk corky patches scored moderate values The highdiversity values for the characters showed a wide variabilityamong genotypes According to [54] this index is also usedin genetic resource studies as a convenient measure of bothallelic richness and allelic evenness when using geneticdata
The highest diversity computed was on presence orabsence of bud groove with a diversity index of 100indicating that this character is very variable in the localsugarcane germplasm On the other hand the descriptorcomputed with the lowest diversity index value (049) wasthe presence or absence of stalk corky cracks indicatingthat this character is selected in the development of thegenotypes In agreement with the present study [55] workingon S officinarum accessions from the world collection ofsugarcane germplasm obtained a low index of 025 and 035for presence or absence of stalk corky patches and cracksrespectively
Their report also showed high diversity index values forinternode alignment growth cracks internode shape budcushion bud extension bud groove and ligule shape Sim-ilarly [56] experimenting on field collections of sugarcaneaccessions in Philippine reported high ShannonndashWeaverdiversity for stalk growth cracks bud cushion stalk corkypatch stalk corky cracks bud extension and auricle outershape Unlike the present study they have reported high indexfor stalk corky cracksMedium index values were reported forstalk internode shape and low index values for ligule shapeand bud shape High variability of sugarcane varieties fordewlap shape was also reported in [57]
The H1015840 pooled across characters by region ranged from079 in Oromiya to 085 in Tigray with an overall aver-age of 081 (Table 8) Most of the regions showed highShannonndashWeaver diversity index The populations that hadthe highest H1015840 were that of Tigray and introduction The H1015840averaged over regions for different characters was found torange from 044 for presence or absence of stalk corky cracksto 094 for relative degree of internode alignment and budgroove with an overall average of 081 (Table 9)
Assessment of the ShannonndashWeaver index also showedsensitivity to both the number of descriptor classes and the
distribution within phenotypic classes in each region Amongregions the mean values for the ShannonndashWeaver index arenot significantly different but differences were found whenindividual characters were considered These observationssuggest that some regions have high diversity for particulartraits while in other regions selective pressures might havereduced the variation to certain genotypes
4 Conclusions
The large number of landraces observed during the cur-rent study demonstrates the existence of diverse geneticresources of sugarcane in Ethiopia The landraces wouldserve to broaden the genetic base of the available sugarcanegermplasm Generally distribution of the overall phenotypicdiversity among sugarcane landraces in Ethiopia is unevenOur results showed that there was a wide range of variationthat existed in the local sugarcane landraces studied both atregional and within region levels Selection and differenti-ation of types might have taken place along a geographicalpattern perhaps associated with climate and use The presentstudy contributes to updating sugarcane descriptors adoptedfrom USDA-ARS as well as Bioversity passport data forthe future collection and evaluation As Ethiopian SugarCorporation is in the course of launching breeding programthe observed substantial variation of landraces would enablesugarcane breeders to design and practice breeding andselection programs to improve the crop
Data Availability
The data used to support the findings of this study areavailable from the corresponding author upon request
Conflicts of Interest
The authors declare that they have no conflicts of interest
Acknowledgments
The authors are grateful for the financial grant of SugarCorporation of Ethiopia Many thanks are due to EthiopianBiodiversity Institute (EBI) for its technical advice duringcollection of the local sugarcane genotypes Sincere thanksare forwarded to Ethiopian smallholder farmers who pre-served sugarcane germplasm and shared their knowledgeand experience Regional Zonal and District (ldquoWeredardquo)Agriculture Bureaus across Ethiopia are acknowledged fortheir support during the study
Supplementary Materials
These supplementary materials contain different tables sup-porting the results of the study These include passport dataof sugarcane germplasm collected and related informationon the materials local sugarcane germplasm collected andused formorphological diversity study introduced sugarcane
16 Advances in Agriculture
Table9Sh
anno
n-WeaverD
iversityIndices(H1015840)a
ndmeanandsta
ndarderror(plusmnS
E)of
land
racesineach
region
basedon
16discretecharacters
Characters
Amha
raBe
nsha
ngul-G
umz
Introd
uced
Oromia
SNNP
Tigray
ELb
Total
Presence
orabsenceo
fbud
cushion(BUDCUSH
ION)
083
095
095
089
082
098
090
088
Relativ
edegreeo
fbud
extension(BUDEX
TEND)
077
079
093
074
072
087
080
078
Relativ
ebud
Shape(BU
DSH
APE
)078
096
089
081
085
078
084
078
Canop
ystructure(C
ANOPY
)089
098
095
081
078
096
089
084
Relativ
eshape
ofdewlap(D
EWLA
PSHAP)
087
094
092
090
082
096
090
086
Relativ
eplant
erectness(ER
ECT)
080
082
050
067
071
088
073
064
Relativ
edegreeo
finterno
dealignm
ent(IN
ALIGN)
096
076
098
097
098
100
094
098
Relativ
einterno
deshape(IN
SHAPE
)086
069
095
077
083
091
083
082
Color
oftheleaves(LE
AFC
OLO
R)087
071
090
064
074
071
076
076
Type
ofou
tera
uricle(A
URI
CLE
OUT)
080
092
087
079
083
093
086
082
Relativ
eshape
ofligule(LIGSH
APE
)086
094
082
085
090
083
087
083
Presence
orabsenceo
fstalkcorkyc
racks(ST
ALK
CORK
C)
044
054
035
055
051
025
044
049
Presence
orabsenceo
fstalkcorkyp
atches(STA
LKCORK
P)066
000
072
079
059
087
060
069
Presence
orabsenceo
fstalkgrow
thcracks
(STA
LKCRA
CK)
066
100
088
062
088
099
084
080
Color
ofthee
xposed
rind(R
INDCOLE
)085
096
089
084
086
076
086
085
Presence
orabsenceo
fbud
groo
ve(BUDGRO
OVE)
095
100
072
100
100
099
094
100
Means
080plusmn0
03
081plusmn0
06
083plusmn0
04
079plusmn0
03
080plusmn0
03
085plusmn0
05
081plusmn0
03
080
b EL=
averageo
fentire
locatio
n
Advances in Agriculture 17
varieties used for morphological diversity study and fre-quency distribution for 16 qualitative characters in sugarcaneby regions in Ethiopia and country of sources of introducedgenotypes (Supplementary Materials)
References
[1] ldquoSpreading the word about clean solutions from sugarcanrdquoAccessed 04072018 httpwwwsugarcaneorg
[2] Central Statistics Agency (CSA) ldquoAgricultural sample surveyArea and production of major crops private peasant holdingsrdquoStatistical Bulletin Addis Ababa Ethiopia 2017
[3] T Esayas Exploration and Collection Characterization GeneticDiversity Analysis and Association of Traits for Yield and YieldComponents of Sugarcane (Saccharumspp) in Ethiopia [MasterThesis] Haramaya University Dire Dawa Ethiopia 2014
[4] J R Harlan Crops and Man American Society of AgronomyCrop Science Society of America Madison WI USA 2ndedition 1992
[5] R W Allard Principles of Plant Breeding JohnWilley and SonsNew York NY USA 2nd edition 1999
[6] B T Roach and J Daniels ldquoA review of the origin and improve-ment of sugarcanerdquo in Proceedings of Coper sugar InternationalSugarcane Workshop pp 1ndash32 1987
[7] P P Patil S S Patil and D U Patel ldquoGenetic diversity in sug-arcane (Saccharum spp Complex)rdquo Journal of Pharmacognosyand Phytochemistry vol 6 no 6 pp 570ndash573 2017
[8] R B Singh B Singh andR K Singh ldquoStudy of genetic diversityof sugarcane (Saccharum) species and commercial varietiesthrough TRAP molecular markersrdquo Indian Journal of PlantPhysiology vol 22 no 3 pp 332ndash338 2017
[9] R W Allard ldquoPopulation structure and sampling methodsrdquo inGenetic Resources in PlantsmdashTheir Exploration and Conserva-tion O H Frankel and E Bennett Eds IBP Handbook no 11pp 97ndash107 FA Davis Company Philadelphia PA USA 1970
[10] D R Marshall and A H D Brown ldquoOptimum samplingstrategies in genetic conservationrdquo inCrop Genetic Resources forToday and Tomorrow O H Frankel and J G Hawkes Eds pp53ndash80 Cambridge University Press 1975
[11] J G Hawkes Crop Genetic Resources Field Collection ManualPudoc Wagnningen Netherland 1980
[12] J G Hawkes The Diversity of Crop Plants Harvard UniversityPress London England 1983
[13] B P K Yerima Manual for Good Laboratory Practice Foodand Agriculture Organization of United Nation Addis AbabaEthiopia 1993
[14] A Walkley and I A Black ldquoAn examination of the degtjareffmethod for determining soil organic matter and a proposedmodification of the chromic acid titrationmethodrdquo Soil Sciencevol 37 no 1 pp 29ndash38 1934
[15] S R Olsen and L E Sommers ldquoPhosphorusrdquo in AgronomyMonograph A L Page et al Ed Methods of soil analysis part2 pp 403ndash430 ASA and SSSAMadisonWI USA 2nd edition1982
[16] R H Bray and L T Kurtz ldquoDetermination of total organic andavailable forms of phosphorus in soilsrdquo Soil Science vol 59 no1 pp 39ndash46 1945
[17] S Sahlemedhin andB Taye ldquoProcedures for soil and plant anal-ysisrdquo Tech Rep 74 National Soil Research Center EthiopianAgricultural Research Organization 2000
[18] F C Stickler S Weaden and AW Pauli ldquoLeaf area determina-tion in grain sorghumrdquo Journal of Agronomy and Crop Sciencevol 53 pp 187-188 1961
[19] Munsell Color ChartThe Munsell Book of Color Matte Collec-tion Munsell Color Chart NewWindsor NY USA 1990
[20] H O Hartley ldquoThe maximum F-ratio as a short cut test forheterogeneity of variancesrdquo Biometrika vol 37 pp 308ndash3121950
[21] L Pecetti P Annicchiarico and A B Damania ldquoBiodiversity ina germplasm collection of durumwheatrdquo Euphytica vol 60 no3 pp 229ndash238 1992
[22] L Pecetti and A B Damania ldquoGeographic variation intetraploid wheat (Triticum turgidum ssp turgidum convardurum) landraces from two provinces in Ethiopiardquo GeneticResources and Crop Evolution vol 43 no 5 pp 395ndash407 1996
[23] S K Jain C O Qualset G M Bhatt and K K Wu ldquoGeo-graphical patterns of phenotypic diversity in a world collectionof durumwheatsrdquoCrop Science vol 15 no 5 pp 700ndash704 1975
[24] K K Nkongolo L Chinthu M Malusi and Z Vokhiwa ldquoPar-ticipatory variety selection and characterization of sorghum(Sorghum bicolour (L) Moench) elite accessions from Malaw-ian gene pool using farmer and breeder knowledgerdquo AfricanJournal of Agricultural Research vol 3 pp 273ndash283 2008
[25] M B Semira ldquoInnovative methods and metrics for agricul-ture and nutrition actions soil quality and human nutritionin Ethiopiardquo Leverhulme Center for Integrative Research onAgriculture and Health (LCIRAH) 2017
[26] I A Khan A Khatri M A Siddiqui G S Nizamani and SRaza ldquoPerformance of promising sugarcane clone for yield andquality traits in different ecological zones of Sindhirdquo PakistanJournal of Botany vol 36 no 1 pp 83ndash92 2004
[27] M F Perera M E Arias D Costilla et al ldquoGenetic diversityassessment and genotype identification in sugarcane based onDNAmarkers andmorphological traitsrdquo Euphytica vol 185 no3 pp 491ndash510 2012
[28] C G Neil M Kay and C C Jack ldquoDiversity among mainlandUSA sugarcane cultivars examined by SSR genotypingrdquo Journalof the American Society of Sugar Cane Technologists vol 29 pp36ndash52 2009
[29] S SchenckMWCrepeauK KWu P HMooreQ Yu andRMing ldquoGenetic diversity and relationships in native HawaiianSaccharum officinarum sugarcanerdquo Journal of Heredity vol 95no 4 pp 327ndash331 2004
[30] R Gebretsadik H Shimelis M D Laing P Tongoona andN Mandefro ldquoA diagnostic appraisal of the sorghum farmingsystem and breeding priorities in Striga infested agro-ecologiesof Ethiopiardquo Agricultural Systems vol 123 pp 54ndash61 2014
[31] L E V Cesar A T Bruzi J A R Nunes et al ldquoGeneticdivergence of sugarcane varieties collected in the region ofLavras Minas Gerais Brazilrdquo Genetics and Molecular Researchvol 14 no 4 pp 14043ndash14049 2015
[32] C Grenier P J Bramel J A Dahlberg et al ldquoSorghums of theSudan analysis of regional diversity and distributionrdquo GeneticResources and Crop Evolution vol 51 no 5 pp 489ndash500 2004
[33] N V Nair R Balakrishnan and T V Sreenivasan ldquoVariabilityfor quantitative traits in exotic hybrid germplasm of sugarcanerdquoGenetic Resources and Crop Evolution vol 45 no 5 pp 459ndash464 1998
[34] R Nagarajan S Alarmelu and R M Shanthi ldquoStudies onvariation in interspecific hybrids of Saccharumrdquo Sugar Tech vol2 no 3 pp 42ndash46 2000
18 Advances in Agriculture
[35] D N Kamat and J R P Singh ldquoVariability in Sugarcane underrain fed conditionrdquo Sugar Technology Short Communicationsvol 3 no 1-2 pp 65ndash67 2001
[36] B S Rishipal A S M Chaudhary and S P Kadian ldquoStudieson character interrelationship in clonal generation of sugarcane(Saccharum complex hybrids)rdquo Indian Sugar vol 47 no 11 pp907ndash911 1998
[37] S Thippeswamy S T Kajjidoni P M Salimath and J V GoudldquoCorrelation and path analysis for cane yield juice quality andtheir component traits in sugarcanerdquo Short Communication vol5 no 1-2 pp 65ndash72 2003
[38] P S Verma S Pal and N K Karma ldquoGenetic variability andcorrelation studies in sugarcanerdquo Indian Sugar vol 49 pp 125ndash128 1999
[39] H A Esechie J W Maranville and W M Ross ldquoRelationshipof stalk morphology and chemical composition to lodgingresistance in sorghumrdquo Crop Science vol 17 no 4 p 609 1977
[40] P K Das B C Jena and N Nayak ldquoCharacter association andpath analysis of sugar yield in sugarcanerdquo Indian Sugar vol 46no 10 pp 805ndash808 1997
[41] K J Sreekumar M Kuriakose M Thomas D Alexander andS Santhakumari ldquoVariability heritability and correlation onstudies on the yield and quality characters of sugarcanerdquo IndianSugar vol 44 no 4 pp 243ndash249 1994
[42] J R Harlan J M de Wet and E G Price ldquoComparativeevolution of cerealsrdquo Evolution vol 27 no 2 pp 311ndash325 1973
[43] A Elings ldquoDurum wheat landraces from syria II patterns ofvariationrdquo Euphytica vol 54 no 3 pp 231ndash243 1991
[44] K Tato ldquoRainfall in Ethiopiardquo Ethiopian Geographical Journalvol 2 pp 28ndash36 1964
[45] A B L Stemler J R Harlan and J M J deWet ldquoThe sorghumsof ethiopiardquo Economic Botany vol 31 no 4 pp 446ndash460 1977
[46] P Annicchiarico L Pecetti and A B Damania ldquoRelationshipsbetween phenotypic variation and climatic factors at collectingsites in durum wheat landracesrdquo Hereditas vol 122 no 2 pp163ndash167 1995
[47] A B Damania L Pecetti C O Qualset and B O HumeidldquoDiversity and geographic distribution of adaptive traits inTriticum turgidum L (durum group) wheat landraces fromTurkeyrdquoGenetic Resources and Crop Evolution vol 43 no 5 pp409ndash422 1996
[48] A Ayana and E Bekele ldquoGeographical patterns of morpho-logical variation in sorghum (Sorghum bicolor (L) Moench)germplasm from Ethiopia and Eritrea quantitative charactersrdquoEuphytica vol 115 no 2 pp 91ndash104 2000
[49] D S Falconer Introduction to Quantitative Genetics JohnWilley and Sons Inc New York NY USA 3rd edition 1993
[50] R S Thorpe ldquoBiochemical analysis of geographical variationand racial affinitiesrdquo Biology Review vol 51 pp 407ndash452 1976
[51] M Farooq ldquoMorphological and agricultural characteristicsof sugarcane in relation to mechanized agriculture at NARCIslamabadrdquo Portu Rico Journal vol 1 pp 51ndash61 1989
[52] M Akhtar N N Elahi and M Ashraf ldquoEvaluation of exoticsugarcane varieties for agronomic characters and productivityrdquoPakistan Journal of Biological Sciences vol 4 no 1 pp 37ndash402001
[53] F R Piscitelli ldquoPrincipals caracteres exomorfologicos de cul-tures de cana de azucarrdquo TUC Iindustraly Agricos vol 7 pp 49ndash57 1994 (Spanish)
[54] A H D Brown and B S Weir ldquoMeasuring genetic variability inplant populationsrdquo in Isozyme in Plant Genetics Breeding S DTanksley and T J Orton Eds part A Elsevier Science 1983
[55] R Balakrishnan N V Nair and T V Sreenivasan ldquoA methodfor establishing a core collection of Saccharum officinarum Lgermplasm based on quantitative-morphological datardquo GeneticResources and Crop Evolution vol 47 no 1 pp 1ndash9 2000
[56] C A Nestor and B Q Reynaldo Sugarcane Variety Improve-ment in Southeast Asia and the Pacific for Enhanced andSustainable Productivity ndash Germplasm and Disease IndexingComponents Institute of Plant Breeding (IPB) College ofAgriculture (CA) UP Los Banos (UPLB) 2003
[57] A Maqbool J Muhammad and A Sagheer ldquoAgronomic traitsand morphological characteristics of some exotic varieties ofsugarcanerdquoPakistan Journal of Agricultural Research vol 19 no4 2006
This is in agreement with previous studies on sugarcanegermplasm [33ndash35]
This is the first study to report information on geo-graphical pattern of variation in Ethiopian local sugarcanegermplasm which has been lacking so far Our results pro-vided experimental evidence on occurrence of geographicalvariation and significant within-region variation where it washigh in the regions such as Amhara Benshangul-Gumz andSNNPR
The overall patterns of similarity or difference betweenregions seemed to depend on environmental factors such asrainfall temperature length of growing season and altitudeWide variability of millable stalk count at harvest singlecane weight and plant height was observed among regionswhere these characters demonstrated high correlation withcane and sugar yield (Table 7) Therefore breeders can utilizeaccessions of regions showing variability for these charactersin selection programs and to design breeding strategies toproduce varieties with best commercial merits
Bivariate Statistics Correlation coefficients worked out onthe 21 quantitative characters are shown for the entire data(Table 7) between regions (Table 8) and within regions(data not shown) Results of the association studies for theentire data showed that cane and sugar yield had highlysignificant positive correlation with most of the quantitativestalk characters and juice quality parameters However theassociation of tiller counts 4 and 5 months after plantingstalk count 10 months after planting millable stalk countat harvest single cane weight and plant height with caneand sugar yield was stronger suggesting these componentcharacters asmain contributing factor to cane and sugar yieldIn fact sugar yield is the product of cane yield and sucrose per-cent cane surely being influenced with components affectingcane yield Similar trend was reported by earlier sugarcaneworkers [36 37] In sugarcane the cane and sugar yields areconsidered to be the complex characters The information onthe phenotypic interrelationship of cane yield and sugar yieldwith their component character per se would be of immensehelp to sugarcane breeders
Stem diameter had significant positive association withboth cane and sugar yield indicating the significance ofthis trait in improving both cane and sugar yields Similarreports have beenmade by earlier sugarcane workers [37 38]Stalk diameter was also positively and significantly correlatedwith plant height It is one of the traits that are related tolodging resistance [39] and the positive associations withheight would help in reducing the chance of lodging as heightincreases Among the physiological attributes included in thestudy leaf area had highly significant positive associationwith cane yield Similar observation was also made in [37]Sucrose percent which is an important juice quality attributehad highly significant positive association with sugar yieldand simultaneously cane yield also had highly significantassociation with sugar yield Based on the magnitude ofcorrelation coefficient values it can be said that cane yield ismuch more important than sucrose percent in determiningthe sugar yield and the same has also been reported by othersugarcane workers [37 40] Similar to observations made in
[37 41] results of the current study showed that all the qualityparameters like sucrose percent pol percent brix percentand purity percent were significantly correlated in positivedirection This indicated any of these juice quality traitscould be considered for selection leading to the simultaneousimprovement in the remaining quality traits
Ten of the 21 characters also showed significant positivecorrelations with altitude of the collection sites (Table 7) Asdescribed in [42] ecological characteristics have influencedthe genotypic constitution of landraces during domesticationand hence a relationship exists between the agroecology inthe collection site and the morphological characteristics ofthe landraces Thus positive correlation between collectionsite variables and plant characteristics would imply that thevariation between accessionsmay be related to agroecologicalvariations among the collection sites [43]
The correlation coefficient between number of internodesand altitude was negative and significant indicating thatother environmental factors (other than altitude) andornonenvironmental factorsmight account for the variation forthis particular character Since temperature decreases withan increase of altitude in Ethiopia [44 45] it is more likelythat temperature has exerted strong selection pressure innumber of internodes in the genotypes Information on therelationships between environmental factors of the collect-ing sites and morphophysiological variation of germplasmcould enhance the understanding of evolutionary adaptivepatterns which could assist germplasm collectors and users[46]
Character associations between region and within regionalso followed similar fashion like that of the entire dataNumber of sprout tiller and stalk counts and sugar qualityparameters showed significant positive correlations with caneand sugar yield
Though not significant and strong the correlation of leaflength and width and leaf area (leaf area = leaf length Xleaf width X 075) with cane and sugar yield was negativein the interregion correlations This kind of relationship wasalso exhibited between these leaf characteristics and yieldcomponents like tiller and stalks counts plant height andsingle cane weight and stalk diameter and sugar qualityparameters This may have been attributed to the variationof temperature with altitude and other environmental factorsacross the regions resulting in such type of relationshipSuch relationship also calls for the need to have enough dataon environmental variables of the collection sites such astemperature and rainfall if germplasm collection is requiredtomeet its objectives effectively and efficiently Unfortunatelydata on such useful environmental variables like temperatureand rainfall are lacking in the passport data of the currentEthiopian sugarcane germplasm collections The significanceof environmental factors of the collection sites as veryimportant determinants in structuring morphophysiolog-ical variations was reported in tetraploid wheat [22 4748]
Association of characters among yield its componentsand other economic traits is important for the interpretationof the patterns of variation and making selection in breed-ing program and combining several desirable attributes It
suggests the advantage of a scheme of selection for morethan one character at a time The correlation between char-acters may arise from linkage or from developmental geneticinteractions with orwithout a purely phenotypic componentIt could arise also due to differential phenotypic plasticityof characters themselves Within the limit of experimentalerror and environmental effects high correlation coefficientsbetween characters may show that the characters sharesome common element of genetic control (ie pleiotropylinkage) between genes or else from independently controlledcharacters responding similarly to geographic variation inselection pressures [49] The interregion correlation coef-ficient between the characters measures the consistency oftheir patterns of regional variation while the intraregioncorrelation coefficient measures the association arising fromgenetic factors but is not affected by regional variation[50] Since this study showed significant positive correlationsintraregionally for some character combinations it wouldseem that common genetic control might be playing a role inbringing about correlations between the various charactersIt appeared that different response to regional variationwas playing a greater role than different genetic controlas shown by the many more significant and moderate tohigh correlation coefficients intraregionally than interregion-ally
322 Qualitative Characters The frequency distribution forthe 16 qualitative characters of germplasm samples by regionsin Ethiopia and of the exotic accessions is shown in Supple-mental Table 5 For the purpose of avoiding redundancy andimproving readability the discrete characters are presented bygrouping related traits together
BudCushion andRelativeDegree of BudExtension Evaluationof the presence and absence of bud cushion revealed thatgreat proportion (70) of sugarcane landraces from Ethiopiado not have bud cushion Majority of the sample germplasmin all regions belong to this phenotypic class Approximatelyone-half (53) of the collection have their bud extendedabove the growth ring followed by those (41) having budtouching the growth ring
Relative Bud Shape Among the phenotypic classes for thischaracter tall deltoid was dominant (31) followed by ovate(17) and narrow ovate (12) While local accessions inmost regions had tall deltoid bud shape 50 of the intro-duced accessions belong to two phenotypic classes roundwith central germpore (30) and 20 having pentagonalbud shape Equal proportions (19) of the samples fromBenshangul-Gumz were Ovate and Ovate with emarginatebasal wing
Canopy Structure Relative Plant Erectness Colour of theLeaves and Colour of the Exposed Rind Out of the eightphenotypic classes observed compact tip droopy was themost frequent (35) Open semidroopy and open droopyclasses had fairly equal distribution With regard to relativeplant erectness seventy-five percent of the germplasm werefound to fall in twophenotypic classes erect (41) and almost
erect (34) for the whole region and country Most of theaccessions (90) showed light green or green leaf colourbut 20 from introduced collections were greenish yellowAmong phenotypic classes for colour of the exposed rindbrownish yellow yellow brown and yellowish green were themost frequent with fairly equal distribution
Relative Shape of Dewlap Double crescent had larger fre-quency (22) while squarish descending and flaring typeof dewlap shape had fairly equal proportions Seventeenpercent of germplasm from Tigray region had squarishdeltoid dewlap shape
Relative Degree of Internode Alignment and Internode ShapeForty-four percent of the sample germplasm were slightlyzigzag while there was almost equal frequency for straightand zigzag internode alignment for the entire region andcountry Straight or nearly straight (slightly zigzag) internodealignment of cane stalks is a very important character formechanized farming and postharvest handling in sugarcane[51] Forty percent of the collections were observed havingconcave-convex type of internode shape while 29 wascylindrical and 14 conoidal Other classes had fairly equalproportion
Type of Auricle and Relative Shape of Ligule An equalproportion of the germplasm totally constituting 84 ofthe collection belong to four phenotypic classes namelythose with no auricle and others with transitional and shortand lanceolate auricles Sixteen percent of the collectionsalso showed equal frequency for the remaining classesBenshangul-Gumz and Tigray regions had each equal 13accessions with falcate type of auricle Sixty-seven percentof the collections belong to three ligule shape classes cres-cent with lozenge (27) broad-crescent (21) and linear-crescent (19) The germplasm had almost equal proportionfor the other phenotypic classes of this character Nineteenpercent of the accessions fromBenshangul-Gumz region haddeltoid shape of ligule
Stalk Corky Cracks Corky Patches Growth Cracks and BudGroove Eighty-nine percent of the germplasm across regionshad no stalk corky cracks with only eleven percent exhibitingthis character All the germplasm in Gambella Harari andSomali had no stalk corky cracks On the other hand eighty-one percent of the germplasm for the whole region showedthe presence of stalk corky patches All the local germplasmin Benshangul-Gumz Gambella and Tigray had stalk corkypatches whereas 75 of the germplasm from Harari Region67 from Somali Region and 24 from Oromia Regionhad no stalk corky patches In the entire region 75 ofthe germplasm had no stalk growth cracks with only 25showing this character Sixty-three percent of the germplasmwere found to have bud groove Howevermost of the samplesin Harari (75) Amhara (63) and Oromia (53) Regionshad no bud groove
The results of this study indicated the wide distributionof phenotypic classes for the characters considered whichindicates the existence of different races and combination
Advances in Agriculture 15
of races in local genotypes of different regions in EthiopiaThis diversity of germplasmbased onphenotypicmarkers canhelp sugarcane breeders in identifying the identity of geno-types as well as maintaining genetic diversity Differencesin morphological characters of different sugarcane varietieshave been reported [52] Piscitelli [53] showed importantexomorphological qualitative characters of sugarcane varietythat are not influenced by environmental factors and thus canbe used as a selection tool in any breeding program
The amount of phenotypic diversity estimates based onthe ShannonndashWeaver diversity index (H1015840) are shown inTable 9 The 16 characters differed in their distribution aswell as the amount of variation Individual traits showed adifferent pattern of variation among accessions Estimatesof H1015840 for individual characters varied from 049 for stalkcorky cracks to 100 for bud groove with an overall meanof 080 Most of the characters were highly polymorphicwhereas relative plant erectness and presence or absenceof stalk corky patches scored moderate values The highdiversity values for the characters showed a wide variabilityamong genotypes According to [54] this index is also usedin genetic resource studies as a convenient measure of bothallelic richness and allelic evenness when using geneticdata
The highest diversity computed was on presence orabsence of bud groove with a diversity index of 100indicating that this character is very variable in the localsugarcane germplasm On the other hand the descriptorcomputed with the lowest diversity index value (049) wasthe presence or absence of stalk corky cracks indicatingthat this character is selected in the development of thegenotypes In agreement with the present study [55] workingon S officinarum accessions from the world collection ofsugarcane germplasm obtained a low index of 025 and 035for presence or absence of stalk corky patches and cracksrespectively
Their report also showed high diversity index values forinternode alignment growth cracks internode shape budcushion bud extension bud groove and ligule shape Sim-ilarly [56] experimenting on field collections of sugarcaneaccessions in Philippine reported high ShannonndashWeaverdiversity for stalk growth cracks bud cushion stalk corkypatch stalk corky cracks bud extension and auricle outershape Unlike the present study they have reported high indexfor stalk corky cracksMedium index values were reported forstalk internode shape and low index values for ligule shapeand bud shape High variability of sugarcane varieties fordewlap shape was also reported in [57]
The H1015840 pooled across characters by region ranged from079 in Oromiya to 085 in Tigray with an overall aver-age of 081 (Table 8) Most of the regions showed highShannonndashWeaver diversity index The populations that hadthe highest H1015840 were that of Tigray and introduction The H1015840averaged over regions for different characters was found torange from 044 for presence or absence of stalk corky cracksto 094 for relative degree of internode alignment and budgroove with an overall average of 081 (Table 9)
Assessment of the ShannonndashWeaver index also showedsensitivity to both the number of descriptor classes and the
distribution within phenotypic classes in each region Amongregions the mean values for the ShannonndashWeaver index arenot significantly different but differences were found whenindividual characters were considered These observationssuggest that some regions have high diversity for particulartraits while in other regions selective pressures might havereduced the variation to certain genotypes
4 Conclusions
The large number of landraces observed during the cur-rent study demonstrates the existence of diverse geneticresources of sugarcane in Ethiopia The landraces wouldserve to broaden the genetic base of the available sugarcanegermplasm Generally distribution of the overall phenotypicdiversity among sugarcane landraces in Ethiopia is unevenOur results showed that there was a wide range of variationthat existed in the local sugarcane landraces studied both atregional and within region levels Selection and differenti-ation of types might have taken place along a geographicalpattern perhaps associated with climate and use The presentstudy contributes to updating sugarcane descriptors adoptedfrom USDA-ARS as well as Bioversity passport data forthe future collection and evaluation As Ethiopian SugarCorporation is in the course of launching breeding programthe observed substantial variation of landraces would enablesugarcane breeders to design and practice breeding andselection programs to improve the crop
Data Availability
The data used to support the findings of this study areavailable from the corresponding author upon request
Conflicts of Interest
The authors declare that they have no conflicts of interest
Acknowledgments
The authors are grateful for the financial grant of SugarCorporation of Ethiopia Many thanks are due to EthiopianBiodiversity Institute (EBI) for its technical advice duringcollection of the local sugarcane genotypes Sincere thanksare forwarded to Ethiopian smallholder farmers who pre-served sugarcane germplasm and shared their knowledgeand experience Regional Zonal and District (ldquoWeredardquo)Agriculture Bureaus across Ethiopia are acknowledged fortheir support during the study
Supplementary Materials
These supplementary materials contain different tables sup-porting the results of the study These include passport dataof sugarcane germplasm collected and related informationon the materials local sugarcane germplasm collected andused formorphological diversity study introduced sugarcane
16 Advances in Agriculture
Table9Sh
anno
n-WeaverD
iversityIndices(H1015840)a
ndmeanandsta
ndarderror(plusmnS
E)of
land
racesineach
region
basedon
16discretecharacters
Characters
Amha
raBe
nsha
ngul-G
umz
Introd
uced
Oromia
SNNP
Tigray
ELb
Total
Presence
orabsenceo
fbud
cushion(BUDCUSH
ION)
083
095
095
089
082
098
090
088
Relativ
edegreeo
fbud
extension(BUDEX
TEND)
077
079
093
074
072
087
080
078
Relativ
ebud
Shape(BU
DSH
APE
)078
096
089
081
085
078
084
078
Canop
ystructure(C
ANOPY
)089
098
095
081
078
096
089
084
Relativ
eshape
ofdewlap(D
EWLA
PSHAP)
087
094
092
090
082
096
090
086
Relativ
eplant
erectness(ER
ECT)
080
082
050
067
071
088
073
064
Relativ
edegreeo
finterno
dealignm
ent(IN
ALIGN)
096
076
098
097
098
100
094
098
Relativ
einterno
deshape(IN
SHAPE
)086
069
095
077
083
091
083
082
Color
oftheleaves(LE
AFC
OLO
R)087
071
090
064
074
071
076
076
Type
ofou
tera
uricle(A
URI
CLE
OUT)
080
092
087
079
083
093
086
082
Relativ
eshape
ofligule(LIGSH
APE
)086
094
082
085
090
083
087
083
Presence
orabsenceo
fstalkcorkyc
racks(ST
ALK
CORK
C)
044
054
035
055
051
025
044
049
Presence
orabsenceo
fstalkcorkyp
atches(STA
LKCORK
P)066
000
072
079
059
087
060
069
Presence
orabsenceo
fstalkgrow
thcracks
(STA
LKCRA
CK)
066
100
088
062
088
099
084
080
Color
ofthee
xposed
rind(R
INDCOLE
)085
096
089
084
086
076
086
085
Presence
orabsenceo
fbud
groo
ve(BUDGRO
OVE)
095
100
072
100
100
099
094
100
Means
080plusmn0
03
081plusmn0
06
083plusmn0
04
079plusmn0
03
080plusmn0
03
085plusmn0
05
081plusmn0
03
080
b EL=
averageo
fentire
locatio
n
Advances in Agriculture 17
varieties used for morphological diversity study and fre-quency distribution for 16 qualitative characters in sugarcaneby regions in Ethiopia and country of sources of introducedgenotypes (Supplementary Materials)
References
[1] ldquoSpreading the word about clean solutions from sugarcanrdquoAccessed 04072018 httpwwwsugarcaneorg
[2] Central Statistics Agency (CSA) ldquoAgricultural sample surveyArea and production of major crops private peasant holdingsrdquoStatistical Bulletin Addis Ababa Ethiopia 2017
[3] T Esayas Exploration and Collection Characterization GeneticDiversity Analysis and Association of Traits for Yield and YieldComponents of Sugarcane (Saccharumspp) in Ethiopia [MasterThesis] Haramaya University Dire Dawa Ethiopia 2014
[4] J R Harlan Crops and Man American Society of AgronomyCrop Science Society of America Madison WI USA 2ndedition 1992
[5] R W Allard Principles of Plant Breeding JohnWilley and SonsNew York NY USA 2nd edition 1999
[6] B T Roach and J Daniels ldquoA review of the origin and improve-ment of sugarcanerdquo in Proceedings of Coper sugar InternationalSugarcane Workshop pp 1ndash32 1987
[7] P P Patil S S Patil and D U Patel ldquoGenetic diversity in sug-arcane (Saccharum spp Complex)rdquo Journal of Pharmacognosyand Phytochemistry vol 6 no 6 pp 570ndash573 2017
[8] R B Singh B Singh andR K Singh ldquoStudy of genetic diversityof sugarcane (Saccharum) species and commercial varietiesthrough TRAP molecular markersrdquo Indian Journal of PlantPhysiology vol 22 no 3 pp 332ndash338 2017
[9] R W Allard ldquoPopulation structure and sampling methodsrdquo inGenetic Resources in PlantsmdashTheir Exploration and Conserva-tion O H Frankel and E Bennett Eds IBP Handbook no 11pp 97ndash107 FA Davis Company Philadelphia PA USA 1970
[10] D R Marshall and A H D Brown ldquoOptimum samplingstrategies in genetic conservationrdquo inCrop Genetic Resources forToday and Tomorrow O H Frankel and J G Hawkes Eds pp53ndash80 Cambridge University Press 1975
[11] J G Hawkes Crop Genetic Resources Field Collection ManualPudoc Wagnningen Netherland 1980
[12] J G Hawkes The Diversity of Crop Plants Harvard UniversityPress London England 1983
[13] B P K Yerima Manual for Good Laboratory Practice Foodand Agriculture Organization of United Nation Addis AbabaEthiopia 1993
[14] A Walkley and I A Black ldquoAn examination of the degtjareffmethod for determining soil organic matter and a proposedmodification of the chromic acid titrationmethodrdquo Soil Sciencevol 37 no 1 pp 29ndash38 1934
[15] S R Olsen and L E Sommers ldquoPhosphorusrdquo in AgronomyMonograph A L Page et al Ed Methods of soil analysis part2 pp 403ndash430 ASA and SSSAMadisonWI USA 2nd edition1982
[16] R H Bray and L T Kurtz ldquoDetermination of total organic andavailable forms of phosphorus in soilsrdquo Soil Science vol 59 no1 pp 39ndash46 1945
[17] S Sahlemedhin andB Taye ldquoProcedures for soil and plant anal-ysisrdquo Tech Rep 74 National Soil Research Center EthiopianAgricultural Research Organization 2000
[18] F C Stickler S Weaden and AW Pauli ldquoLeaf area determina-tion in grain sorghumrdquo Journal of Agronomy and Crop Sciencevol 53 pp 187-188 1961
[19] Munsell Color ChartThe Munsell Book of Color Matte Collec-tion Munsell Color Chart NewWindsor NY USA 1990
[20] H O Hartley ldquoThe maximum F-ratio as a short cut test forheterogeneity of variancesrdquo Biometrika vol 37 pp 308ndash3121950
[21] L Pecetti P Annicchiarico and A B Damania ldquoBiodiversity ina germplasm collection of durumwheatrdquo Euphytica vol 60 no3 pp 229ndash238 1992
[22] L Pecetti and A B Damania ldquoGeographic variation intetraploid wheat (Triticum turgidum ssp turgidum convardurum) landraces from two provinces in Ethiopiardquo GeneticResources and Crop Evolution vol 43 no 5 pp 395ndash407 1996
[23] S K Jain C O Qualset G M Bhatt and K K Wu ldquoGeo-graphical patterns of phenotypic diversity in a world collectionof durumwheatsrdquoCrop Science vol 15 no 5 pp 700ndash704 1975
[24] K K Nkongolo L Chinthu M Malusi and Z Vokhiwa ldquoPar-ticipatory variety selection and characterization of sorghum(Sorghum bicolour (L) Moench) elite accessions from Malaw-ian gene pool using farmer and breeder knowledgerdquo AfricanJournal of Agricultural Research vol 3 pp 273ndash283 2008
[25] M B Semira ldquoInnovative methods and metrics for agricul-ture and nutrition actions soil quality and human nutritionin Ethiopiardquo Leverhulme Center for Integrative Research onAgriculture and Health (LCIRAH) 2017
[26] I A Khan A Khatri M A Siddiqui G S Nizamani and SRaza ldquoPerformance of promising sugarcane clone for yield andquality traits in different ecological zones of Sindhirdquo PakistanJournal of Botany vol 36 no 1 pp 83ndash92 2004
[27] M F Perera M E Arias D Costilla et al ldquoGenetic diversityassessment and genotype identification in sugarcane based onDNAmarkers andmorphological traitsrdquo Euphytica vol 185 no3 pp 491ndash510 2012
[28] C G Neil M Kay and C C Jack ldquoDiversity among mainlandUSA sugarcane cultivars examined by SSR genotypingrdquo Journalof the American Society of Sugar Cane Technologists vol 29 pp36ndash52 2009
[29] S SchenckMWCrepeauK KWu P HMooreQ Yu andRMing ldquoGenetic diversity and relationships in native HawaiianSaccharum officinarum sugarcanerdquo Journal of Heredity vol 95no 4 pp 327ndash331 2004
[30] R Gebretsadik H Shimelis M D Laing P Tongoona andN Mandefro ldquoA diagnostic appraisal of the sorghum farmingsystem and breeding priorities in Striga infested agro-ecologiesof Ethiopiardquo Agricultural Systems vol 123 pp 54ndash61 2014
[31] L E V Cesar A T Bruzi J A R Nunes et al ldquoGeneticdivergence of sugarcane varieties collected in the region ofLavras Minas Gerais Brazilrdquo Genetics and Molecular Researchvol 14 no 4 pp 14043ndash14049 2015
[32] C Grenier P J Bramel J A Dahlberg et al ldquoSorghums of theSudan analysis of regional diversity and distributionrdquo GeneticResources and Crop Evolution vol 51 no 5 pp 489ndash500 2004
[33] N V Nair R Balakrishnan and T V Sreenivasan ldquoVariabilityfor quantitative traits in exotic hybrid germplasm of sugarcanerdquoGenetic Resources and Crop Evolution vol 45 no 5 pp 459ndash464 1998
[34] R Nagarajan S Alarmelu and R M Shanthi ldquoStudies onvariation in interspecific hybrids of Saccharumrdquo Sugar Tech vol2 no 3 pp 42ndash46 2000
18 Advances in Agriculture
[35] D N Kamat and J R P Singh ldquoVariability in Sugarcane underrain fed conditionrdquo Sugar Technology Short Communicationsvol 3 no 1-2 pp 65ndash67 2001
[36] B S Rishipal A S M Chaudhary and S P Kadian ldquoStudieson character interrelationship in clonal generation of sugarcane(Saccharum complex hybrids)rdquo Indian Sugar vol 47 no 11 pp907ndash911 1998
[37] S Thippeswamy S T Kajjidoni P M Salimath and J V GoudldquoCorrelation and path analysis for cane yield juice quality andtheir component traits in sugarcanerdquo Short Communication vol5 no 1-2 pp 65ndash72 2003
[38] P S Verma S Pal and N K Karma ldquoGenetic variability andcorrelation studies in sugarcanerdquo Indian Sugar vol 49 pp 125ndash128 1999
[39] H A Esechie J W Maranville and W M Ross ldquoRelationshipof stalk morphology and chemical composition to lodgingresistance in sorghumrdquo Crop Science vol 17 no 4 p 609 1977
[40] P K Das B C Jena and N Nayak ldquoCharacter association andpath analysis of sugar yield in sugarcanerdquo Indian Sugar vol 46no 10 pp 805ndash808 1997
[41] K J Sreekumar M Kuriakose M Thomas D Alexander andS Santhakumari ldquoVariability heritability and correlation onstudies on the yield and quality characters of sugarcanerdquo IndianSugar vol 44 no 4 pp 243ndash249 1994
[42] J R Harlan J M de Wet and E G Price ldquoComparativeevolution of cerealsrdquo Evolution vol 27 no 2 pp 311ndash325 1973
[43] A Elings ldquoDurum wheat landraces from syria II patterns ofvariationrdquo Euphytica vol 54 no 3 pp 231ndash243 1991
[44] K Tato ldquoRainfall in Ethiopiardquo Ethiopian Geographical Journalvol 2 pp 28ndash36 1964
[45] A B L Stemler J R Harlan and J M J deWet ldquoThe sorghumsof ethiopiardquo Economic Botany vol 31 no 4 pp 446ndash460 1977
[46] P Annicchiarico L Pecetti and A B Damania ldquoRelationshipsbetween phenotypic variation and climatic factors at collectingsites in durum wheat landracesrdquo Hereditas vol 122 no 2 pp163ndash167 1995
[47] A B Damania L Pecetti C O Qualset and B O HumeidldquoDiversity and geographic distribution of adaptive traits inTriticum turgidum L (durum group) wheat landraces fromTurkeyrdquoGenetic Resources and Crop Evolution vol 43 no 5 pp409ndash422 1996
[48] A Ayana and E Bekele ldquoGeographical patterns of morpho-logical variation in sorghum (Sorghum bicolor (L) Moench)germplasm from Ethiopia and Eritrea quantitative charactersrdquoEuphytica vol 115 no 2 pp 91ndash104 2000
[49] D S Falconer Introduction to Quantitative Genetics JohnWilley and Sons Inc New York NY USA 3rd edition 1993
[50] R S Thorpe ldquoBiochemical analysis of geographical variationand racial affinitiesrdquo Biology Review vol 51 pp 407ndash452 1976
[51] M Farooq ldquoMorphological and agricultural characteristicsof sugarcane in relation to mechanized agriculture at NARCIslamabadrdquo Portu Rico Journal vol 1 pp 51ndash61 1989
[52] M Akhtar N N Elahi and M Ashraf ldquoEvaluation of exoticsugarcane varieties for agronomic characters and productivityrdquoPakistan Journal of Biological Sciences vol 4 no 1 pp 37ndash402001
[53] F R Piscitelli ldquoPrincipals caracteres exomorfologicos de cul-tures de cana de azucarrdquo TUC Iindustraly Agricos vol 7 pp 49ndash57 1994 (Spanish)
[54] A H D Brown and B S Weir ldquoMeasuring genetic variability inplant populationsrdquo in Isozyme in Plant Genetics Breeding S DTanksley and T J Orton Eds part A Elsevier Science 1983
[55] R Balakrishnan N V Nair and T V Sreenivasan ldquoA methodfor establishing a core collection of Saccharum officinarum Lgermplasm based on quantitative-morphological datardquo GeneticResources and Crop Evolution vol 47 no 1 pp 1ndash9 2000
[56] C A Nestor and B Q Reynaldo Sugarcane Variety Improve-ment in Southeast Asia and the Pacific for Enhanced andSustainable Productivity ndash Germplasm and Disease IndexingComponents Institute of Plant Breeding (IPB) College ofAgriculture (CA) UP Los Banos (UPLB) 2003
[57] A Maqbool J Muhammad and A Sagheer ldquoAgronomic traitsand morphological characteristics of some exotic varieties ofsugarcanerdquoPakistan Journal of Agricultural Research vol 19 no4 2006
This is in agreement with previous studies on sugarcanegermplasm [33ndash35]
This is the first study to report information on geo-graphical pattern of variation in Ethiopian local sugarcanegermplasm which has been lacking so far Our results pro-vided experimental evidence on occurrence of geographicalvariation and significant within-region variation where it washigh in the regions such as Amhara Benshangul-Gumz andSNNPR
The overall patterns of similarity or difference betweenregions seemed to depend on environmental factors such asrainfall temperature length of growing season and altitudeWide variability of millable stalk count at harvest singlecane weight and plant height was observed among regionswhere these characters demonstrated high correlation withcane and sugar yield (Table 7) Therefore breeders can utilizeaccessions of regions showing variability for these charactersin selection programs and to design breeding strategies toproduce varieties with best commercial merits
Bivariate Statistics Correlation coefficients worked out onthe 21 quantitative characters are shown for the entire data(Table 7) between regions (Table 8) and within regions(data not shown) Results of the association studies for theentire data showed that cane and sugar yield had highlysignificant positive correlation with most of the quantitativestalk characters and juice quality parameters However theassociation of tiller counts 4 and 5 months after plantingstalk count 10 months after planting millable stalk countat harvest single cane weight and plant height with caneand sugar yield was stronger suggesting these componentcharacters asmain contributing factor to cane and sugar yieldIn fact sugar yield is the product of cane yield and sucrose per-cent cane surely being influenced with components affectingcane yield Similar trend was reported by earlier sugarcaneworkers [36 37] In sugarcane the cane and sugar yields areconsidered to be the complex characters The information onthe phenotypic interrelationship of cane yield and sugar yieldwith their component character per se would be of immensehelp to sugarcane breeders
Stem diameter had significant positive association withboth cane and sugar yield indicating the significance ofthis trait in improving both cane and sugar yields Similarreports have beenmade by earlier sugarcane workers [37 38]Stalk diameter was also positively and significantly correlatedwith plant height It is one of the traits that are related tolodging resistance [39] and the positive associations withheight would help in reducing the chance of lodging as heightincreases Among the physiological attributes included in thestudy leaf area had highly significant positive associationwith cane yield Similar observation was also made in [37]Sucrose percent which is an important juice quality attributehad highly significant positive association with sugar yieldand simultaneously cane yield also had highly significantassociation with sugar yield Based on the magnitude ofcorrelation coefficient values it can be said that cane yield ismuch more important than sucrose percent in determiningthe sugar yield and the same has also been reported by othersugarcane workers [37 40] Similar to observations made in
[37 41] results of the current study showed that all the qualityparameters like sucrose percent pol percent brix percentand purity percent were significantly correlated in positivedirection This indicated any of these juice quality traitscould be considered for selection leading to the simultaneousimprovement in the remaining quality traits
Ten of the 21 characters also showed significant positivecorrelations with altitude of the collection sites (Table 7) Asdescribed in [42] ecological characteristics have influencedthe genotypic constitution of landraces during domesticationand hence a relationship exists between the agroecology inthe collection site and the morphological characteristics ofthe landraces Thus positive correlation between collectionsite variables and plant characteristics would imply that thevariation between accessionsmay be related to agroecologicalvariations among the collection sites [43]
The correlation coefficient between number of internodesand altitude was negative and significant indicating thatother environmental factors (other than altitude) andornonenvironmental factorsmight account for the variation forthis particular character Since temperature decreases withan increase of altitude in Ethiopia [44 45] it is more likelythat temperature has exerted strong selection pressure innumber of internodes in the genotypes Information on therelationships between environmental factors of the collect-ing sites and morphophysiological variation of germplasmcould enhance the understanding of evolutionary adaptivepatterns which could assist germplasm collectors and users[46]
Character associations between region and within regionalso followed similar fashion like that of the entire dataNumber of sprout tiller and stalk counts and sugar qualityparameters showed significant positive correlations with caneand sugar yield
Though not significant and strong the correlation of leaflength and width and leaf area (leaf area = leaf length Xleaf width X 075) with cane and sugar yield was negativein the interregion correlations This kind of relationship wasalso exhibited between these leaf characteristics and yieldcomponents like tiller and stalks counts plant height andsingle cane weight and stalk diameter and sugar qualityparameters This may have been attributed to the variationof temperature with altitude and other environmental factorsacross the regions resulting in such type of relationshipSuch relationship also calls for the need to have enough dataon environmental variables of the collection sites such astemperature and rainfall if germplasm collection is requiredtomeet its objectives effectively and efficiently Unfortunatelydata on such useful environmental variables like temperatureand rainfall are lacking in the passport data of the currentEthiopian sugarcane germplasm collections The significanceof environmental factors of the collection sites as veryimportant determinants in structuring morphophysiolog-ical variations was reported in tetraploid wheat [22 4748]
Association of characters among yield its componentsand other economic traits is important for the interpretationof the patterns of variation and making selection in breed-ing program and combining several desirable attributes It
suggests the advantage of a scheme of selection for morethan one character at a time The correlation between char-acters may arise from linkage or from developmental geneticinteractions with orwithout a purely phenotypic componentIt could arise also due to differential phenotypic plasticityof characters themselves Within the limit of experimentalerror and environmental effects high correlation coefficientsbetween characters may show that the characters sharesome common element of genetic control (ie pleiotropylinkage) between genes or else from independently controlledcharacters responding similarly to geographic variation inselection pressures [49] The interregion correlation coef-ficient between the characters measures the consistency oftheir patterns of regional variation while the intraregioncorrelation coefficient measures the association arising fromgenetic factors but is not affected by regional variation[50] Since this study showed significant positive correlationsintraregionally for some character combinations it wouldseem that common genetic control might be playing a role inbringing about correlations between the various charactersIt appeared that different response to regional variationwas playing a greater role than different genetic controlas shown by the many more significant and moderate tohigh correlation coefficients intraregionally than interregion-ally
322 Qualitative Characters The frequency distribution forthe 16 qualitative characters of germplasm samples by regionsin Ethiopia and of the exotic accessions is shown in Supple-mental Table 5 For the purpose of avoiding redundancy andimproving readability the discrete characters are presented bygrouping related traits together
BudCushion andRelativeDegree of BudExtension Evaluationof the presence and absence of bud cushion revealed thatgreat proportion (70) of sugarcane landraces from Ethiopiado not have bud cushion Majority of the sample germplasmin all regions belong to this phenotypic class Approximatelyone-half (53) of the collection have their bud extendedabove the growth ring followed by those (41) having budtouching the growth ring
Relative Bud Shape Among the phenotypic classes for thischaracter tall deltoid was dominant (31) followed by ovate(17) and narrow ovate (12) While local accessions inmost regions had tall deltoid bud shape 50 of the intro-duced accessions belong to two phenotypic classes roundwith central germpore (30) and 20 having pentagonalbud shape Equal proportions (19) of the samples fromBenshangul-Gumz were Ovate and Ovate with emarginatebasal wing
Canopy Structure Relative Plant Erectness Colour of theLeaves and Colour of the Exposed Rind Out of the eightphenotypic classes observed compact tip droopy was themost frequent (35) Open semidroopy and open droopyclasses had fairly equal distribution With regard to relativeplant erectness seventy-five percent of the germplasm werefound to fall in twophenotypic classes erect (41) and almost
erect (34) for the whole region and country Most of theaccessions (90) showed light green or green leaf colourbut 20 from introduced collections were greenish yellowAmong phenotypic classes for colour of the exposed rindbrownish yellow yellow brown and yellowish green were themost frequent with fairly equal distribution
Relative Shape of Dewlap Double crescent had larger fre-quency (22) while squarish descending and flaring typeof dewlap shape had fairly equal proportions Seventeenpercent of germplasm from Tigray region had squarishdeltoid dewlap shape
Relative Degree of Internode Alignment and Internode ShapeForty-four percent of the sample germplasm were slightlyzigzag while there was almost equal frequency for straightand zigzag internode alignment for the entire region andcountry Straight or nearly straight (slightly zigzag) internodealignment of cane stalks is a very important character formechanized farming and postharvest handling in sugarcane[51] Forty percent of the collections were observed havingconcave-convex type of internode shape while 29 wascylindrical and 14 conoidal Other classes had fairly equalproportion
Type of Auricle and Relative Shape of Ligule An equalproportion of the germplasm totally constituting 84 ofthe collection belong to four phenotypic classes namelythose with no auricle and others with transitional and shortand lanceolate auricles Sixteen percent of the collectionsalso showed equal frequency for the remaining classesBenshangul-Gumz and Tigray regions had each equal 13accessions with falcate type of auricle Sixty-seven percentof the collections belong to three ligule shape classes cres-cent with lozenge (27) broad-crescent (21) and linear-crescent (19) The germplasm had almost equal proportionfor the other phenotypic classes of this character Nineteenpercent of the accessions fromBenshangul-Gumz region haddeltoid shape of ligule
Stalk Corky Cracks Corky Patches Growth Cracks and BudGroove Eighty-nine percent of the germplasm across regionshad no stalk corky cracks with only eleven percent exhibitingthis character All the germplasm in Gambella Harari andSomali had no stalk corky cracks On the other hand eighty-one percent of the germplasm for the whole region showedthe presence of stalk corky patches All the local germplasmin Benshangul-Gumz Gambella and Tigray had stalk corkypatches whereas 75 of the germplasm from Harari Region67 from Somali Region and 24 from Oromia Regionhad no stalk corky patches In the entire region 75 ofthe germplasm had no stalk growth cracks with only 25showing this character Sixty-three percent of the germplasmwere found to have bud groove Howevermost of the samplesin Harari (75) Amhara (63) and Oromia (53) Regionshad no bud groove
The results of this study indicated the wide distributionof phenotypic classes for the characters considered whichindicates the existence of different races and combination
Advances in Agriculture 15
of races in local genotypes of different regions in EthiopiaThis diversity of germplasmbased onphenotypicmarkers canhelp sugarcane breeders in identifying the identity of geno-types as well as maintaining genetic diversity Differencesin morphological characters of different sugarcane varietieshave been reported [52] Piscitelli [53] showed importantexomorphological qualitative characters of sugarcane varietythat are not influenced by environmental factors and thus canbe used as a selection tool in any breeding program
The amount of phenotypic diversity estimates based onthe ShannonndashWeaver diversity index (H1015840) are shown inTable 9 The 16 characters differed in their distribution aswell as the amount of variation Individual traits showed adifferent pattern of variation among accessions Estimatesof H1015840 for individual characters varied from 049 for stalkcorky cracks to 100 for bud groove with an overall meanof 080 Most of the characters were highly polymorphicwhereas relative plant erectness and presence or absenceof stalk corky patches scored moderate values The highdiversity values for the characters showed a wide variabilityamong genotypes According to [54] this index is also usedin genetic resource studies as a convenient measure of bothallelic richness and allelic evenness when using geneticdata
The highest diversity computed was on presence orabsence of bud groove with a diversity index of 100indicating that this character is very variable in the localsugarcane germplasm On the other hand the descriptorcomputed with the lowest diversity index value (049) wasthe presence or absence of stalk corky cracks indicatingthat this character is selected in the development of thegenotypes In agreement with the present study [55] workingon S officinarum accessions from the world collection ofsugarcane germplasm obtained a low index of 025 and 035for presence or absence of stalk corky patches and cracksrespectively
Their report also showed high diversity index values forinternode alignment growth cracks internode shape budcushion bud extension bud groove and ligule shape Sim-ilarly [56] experimenting on field collections of sugarcaneaccessions in Philippine reported high ShannonndashWeaverdiversity for stalk growth cracks bud cushion stalk corkypatch stalk corky cracks bud extension and auricle outershape Unlike the present study they have reported high indexfor stalk corky cracksMedium index values were reported forstalk internode shape and low index values for ligule shapeand bud shape High variability of sugarcane varieties fordewlap shape was also reported in [57]
The H1015840 pooled across characters by region ranged from079 in Oromiya to 085 in Tigray with an overall aver-age of 081 (Table 8) Most of the regions showed highShannonndashWeaver diversity index The populations that hadthe highest H1015840 were that of Tigray and introduction The H1015840averaged over regions for different characters was found torange from 044 for presence or absence of stalk corky cracksto 094 for relative degree of internode alignment and budgroove with an overall average of 081 (Table 9)
Assessment of the ShannonndashWeaver index also showedsensitivity to both the number of descriptor classes and the
distribution within phenotypic classes in each region Amongregions the mean values for the ShannonndashWeaver index arenot significantly different but differences were found whenindividual characters were considered These observationssuggest that some regions have high diversity for particulartraits while in other regions selective pressures might havereduced the variation to certain genotypes
4 Conclusions
The large number of landraces observed during the cur-rent study demonstrates the existence of diverse geneticresources of sugarcane in Ethiopia The landraces wouldserve to broaden the genetic base of the available sugarcanegermplasm Generally distribution of the overall phenotypicdiversity among sugarcane landraces in Ethiopia is unevenOur results showed that there was a wide range of variationthat existed in the local sugarcane landraces studied both atregional and within region levels Selection and differenti-ation of types might have taken place along a geographicalpattern perhaps associated with climate and use The presentstudy contributes to updating sugarcane descriptors adoptedfrom USDA-ARS as well as Bioversity passport data forthe future collection and evaluation As Ethiopian SugarCorporation is in the course of launching breeding programthe observed substantial variation of landraces would enablesugarcane breeders to design and practice breeding andselection programs to improve the crop
Data Availability
The data used to support the findings of this study areavailable from the corresponding author upon request
Conflicts of Interest
The authors declare that they have no conflicts of interest
Acknowledgments
The authors are grateful for the financial grant of SugarCorporation of Ethiopia Many thanks are due to EthiopianBiodiversity Institute (EBI) for its technical advice duringcollection of the local sugarcane genotypes Sincere thanksare forwarded to Ethiopian smallholder farmers who pre-served sugarcane germplasm and shared their knowledgeand experience Regional Zonal and District (ldquoWeredardquo)Agriculture Bureaus across Ethiopia are acknowledged fortheir support during the study
Supplementary Materials
These supplementary materials contain different tables sup-porting the results of the study These include passport dataof sugarcane germplasm collected and related informationon the materials local sugarcane germplasm collected andused formorphological diversity study introduced sugarcane
16 Advances in Agriculture
Table9Sh
anno
n-WeaverD
iversityIndices(H1015840)a
ndmeanandsta
ndarderror(plusmnS
E)of
land
racesineach
region
basedon
16discretecharacters
Characters
Amha
raBe
nsha
ngul-G
umz
Introd
uced
Oromia
SNNP
Tigray
ELb
Total
Presence
orabsenceo
fbud
cushion(BUDCUSH
ION)
083
095
095
089
082
098
090
088
Relativ
edegreeo
fbud
extension(BUDEX
TEND)
077
079
093
074
072
087
080
078
Relativ
ebud
Shape(BU
DSH
APE
)078
096
089
081
085
078
084
078
Canop
ystructure(C
ANOPY
)089
098
095
081
078
096
089
084
Relativ
eshape
ofdewlap(D
EWLA
PSHAP)
087
094
092
090
082
096
090
086
Relativ
eplant
erectness(ER
ECT)
080
082
050
067
071
088
073
064
Relativ
edegreeo
finterno
dealignm
ent(IN
ALIGN)
096
076
098
097
098
100
094
098
Relativ
einterno
deshape(IN
SHAPE
)086
069
095
077
083
091
083
082
Color
oftheleaves(LE
AFC
OLO
R)087
071
090
064
074
071
076
076
Type
ofou
tera
uricle(A
URI
CLE
OUT)
080
092
087
079
083
093
086
082
Relativ
eshape
ofligule(LIGSH
APE
)086
094
082
085
090
083
087
083
Presence
orabsenceo
fstalkcorkyc
racks(ST
ALK
CORK
C)
044
054
035
055
051
025
044
049
Presence
orabsenceo
fstalkcorkyp
atches(STA
LKCORK
P)066
000
072
079
059
087
060
069
Presence
orabsenceo
fstalkgrow
thcracks
(STA
LKCRA
CK)
066
100
088
062
088
099
084
080
Color
ofthee
xposed
rind(R
INDCOLE
)085
096
089
084
086
076
086
085
Presence
orabsenceo
fbud
groo
ve(BUDGRO
OVE)
095
100
072
100
100
099
094
100
Means
080plusmn0
03
081plusmn0
06
083plusmn0
04
079plusmn0
03
080plusmn0
03
085plusmn0
05
081plusmn0
03
080
b EL=
averageo
fentire
locatio
n
Advances in Agriculture 17
varieties used for morphological diversity study and fre-quency distribution for 16 qualitative characters in sugarcaneby regions in Ethiopia and country of sources of introducedgenotypes (Supplementary Materials)
References
[1] ldquoSpreading the word about clean solutions from sugarcanrdquoAccessed 04072018 httpwwwsugarcaneorg
[2] Central Statistics Agency (CSA) ldquoAgricultural sample surveyArea and production of major crops private peasant holdingsrdquoStatistical Bulletin Addis Ababa Ethiopia 2017
[3] T Esayas Exploration and Collection Characterization GeneticDiversity Analysis and Association of Traits for Yield and YieldComponents of Sugarcane (Saccharumspp) in Ethiopia [MasterThesis] Haramaya University Dire Dawa Ethiopia 2014
[4] J R Harlan Crops and Man American Society of AgronomyCrop Science Society of America Madison WI USA 2ndedition 1992
[5] R W Allard Principles of Plant Breeding JohnWilley and SonsNew York NY USA 2nd edition 1999
[6] B T Roach and J Daniels ldquoA review of the origin and improve-ment of sugarcanerdquo in Proceedings of Coper sugar InternationalSugarcane Workshop pp 1ndash32 1987
[7] P P Patil S S Patil and D U Patel ldquoGenetic diversity in sug-arcane (Saccharum spp Complex)rdquo Journal of Pharmacognosyand Phytochemistry vol 6 no 6 pp 570ndash573 2017
[8] R B Singh B Singh andR K Singh ldquoStudy of genetic diversityof sugarcane (Saccharum) species and commercial varietiesthrough TRAP molecular markersrdquo Indian Journal of PlantPhysiology vol 22 no 3 pp 332ndash338 2017
[9] R W Allard ldquoPopulation structure and sampling methodsrdquo inGenetic Resources in PlantsmdashTheir Exploration and Conserva-tion O H Frankel and E Bennett Eds IBP Handbook no 11pp 97ndash107 FA Davis Company Philadelphia PA USA 1970
[10] D R Marshall and A H D Brown ldquoOptimum samplingstrategies in genetic conservationrdquo inCrop Genetic Resources forToday and Tomorrow O H Frankel and J G Hawkes Eds pp53ndash80 Cambridge University Press 1975
[11] J G Hawkes Crop Genetic Resources Field Collection ManualPudoc Wagnningen Netherland 1980
[12] J G Hawkes The Diversity of Crop Plants Harvard UniversityPress London England 1983
[13] B P K Yerima Manual for Good Laboratory Practice Foodand Agriculture Organization of United Nation Addis AbabaEthiopia 1993
[14] A Walkley and I A Black ldquoAn examination of the degtjareffmethod for determining soil organic matter and a proposedmodification of the chromic acid titrationmethodrdquo Soil Sciencevol 37 no 1 pp 29ndash38 1934
[15] S R Olsen and L E Sommers ldquoPhosphorusrdquo in AgronomyMonograph A L Page et al Ed Methods of soil analysis part2 pp 403ndash430 ASA and SSSAMadisonWI USA 2nd edition1982
[16] R H Bray and L T Kurtz ldquoDetermination of total organic andavailable forms of phosphorus in soilsrdquo Soil Science vol 59 no1 pp 39ndash46 1945
[17] S Sahlemedhin andB Taye ldquoProcedures for soil and plant anal-ysisrdquo Tech Rep 74 National Soil Research Center EthiopianAgricultural Research Organization 2000
[18] F C Stickler S Weaden and AW Pauli ldquoLeaf area determina-tion in grain sorghumrdquo Journal of Agronomy and Crop Sciencevol 53 pp 187-188 1961
[19] Munsell Color ChartThe Munsell Book of Color Matte Collec-tion Munsell Color Chart NewWindsor NY USA 1990
[20] H O Hartley ldquoThe maximum F-ratio as a short cut test forheterogeneity of variancesrdquo Biometrika vol 37 pp 308ndash3121950
[21] L Pecetti P Annicchiarico and A B Damania ldquoBiodiversity ina germplasm collection of durumwheatrdquo Euphytica vol 60 no3 pp 229ndash238 1992
[22] L Pecetti and A B Damania ldquoGeographic variation intetraploid wheat (Triticum turgidum ssp turgidum convardurum) landraces from two provinces in Ethiopiardquo GeneticResources and Crop Evolution vol 43 no 5 pp 395ndash407 1996
[23] S K Jain C O Qualset G M Bhatt and K K Wu ldquoGeo-graphical patterns of phenotypic diversity in a world collectionof durumwheatsrdquoCrop Science vol 15 no 5 pp 700ndash704 1975
[24] K K Nkongolo L Chinthu M Malusi and Z Vokhiwa ldquoPar-ticipatory variety selection and characterization of sorghum(Sorghum bicolour (L) Moench) elite accessions from Malaw-ian gene pool using farmer and breeder knowledgerdquo AfricanJournal of Agricultural Research vol 3 pp 273ndash283 2008
[25] M B Semira ldquoInnovative methods and metrics for agricul-ture and nutrition actions soil quality and human nutritionin Ethiopiardquo Leverhulme Center for Integrative Research onAgriculture and Health (LCIRAH) 2017
[26] I A Khan A Khatri M A Siddiqui G S Nizamani and SRaza ldquoPerformance of promising sugarcane clone for yield andquality traits in different ecological zones of Sindhirdquo PakistanJournal of Botany vol 36 no 1 pp 83ndash92 2004
[27] M F Perera M E Arias D Costilla et al ldquoGenetic diversityassessment and genotype identification in sugarcane based onDNAmarkers andmorphological traitsrdquo Euphytica vol 185 no3 pp 491ndash510 2012
[28] C G Neil M Kay and C C Jack ldquoDiversity among mainlandUSA sugarcane cultivars examined by SSR genotypingrdquo Journalof the American Society of Sugar Cane Technologists vol 29 pp36ndash52 2009
[29] S SchenckMWCrepeauK KWu P HMooreQ Yu andRMing ldquoGenetic diversity and relationships in native HawaiianSaccharum officinarum sugarcanerdquo Journal of Heredity vol 95no 4 pp 327ndash331 2004
[30] R Gebretsadik H Shimelis M D Laing P Tongoona andN Mandefro ldquoA diagnostic appraisal of the sorghum farmingsystem and breeding priorities in Striga infested agro-ecologiesof Ethiopiardquo Agricultural Systems vol 123 pp 54ndash61 2014
[31] L E V Cesar A T Bruzi J A R Nunes et al ldquoGeneticdivergence of sugarcane varieties collected in the region ofLavras Minas Gerais Brazilrdquo Genetics and Molecular Researchvol 14 no 4 pp 14043ndash14049 2015
[32] C Grenier P J Bramel J A Dahlberg et al ldquoSorghums of theSudan analysis of regional diversity and distributionrdquo GeneticResources and Crop Evolution vol 51 no 5 pp 489ndash500 2004
[33] N V Nair R Balakrishnan and T V Sreenivasan ldquoVariabilityfor quantitative traits in exotic hybrid germplasm of sugarcanerdquoGenetic Resources and Crop Evolution vol 45 no 5 pp 459ndash464 1998
[34] R Nagarajan S Alarmelu and R M Shanthi ldquoStudies onvariation in interspecific hybrids of Saccharumrdquo Sugar Tech vol2 no 3 pp 42ndash46 2000
18 Advances in Agriculture
[35] D N Kamat and J R P Singh ldquoVariability in Sugarcane underrain fed conditionrdquo Sugar Technology Short Communicationsvol 3 no 1-2 pp 65ndash67 2001
[36] B S Rishipal A S M Chaudhary and S P Kadian ldquoStudieson character interrelationship in clonal generation of sugarcane(Saccharum complex hybrids)rdquo Indian Sugar vol 47 no 11 pp907ndash911 1998
[37] S Thippeswamy S T Kajjidoni P M Salimath and J V GoudldquoCorrelation and path analysis for cane yield juice quality andtheir component traits in sugarcanerdquo Short Communication vol5 no 1-2 pp 65ndash72 2003
[38] P S Verma S Pal and N K Karma ldquoGenetic variability andcorrelation studies in sugarcanerdquo Indian Sugar vol 49 pp 125ndash128 1999
[39] H A Esechie J W Maranville and W M Ross ldquoRelationshipof stalk morphology and chemical composition to lodgingresistance in sorghumrdquo Crop Science vol 17 no 4 p 609 1977
[40] P K Das B C Jena and N Nayak ldquoCharacter association andpath analysis of sugar yield in sugarcanerdquo Indian Sugar vol 46no 10 pp 805ndash808 1997
[41] K J Sreekumar M Kuriakose M Thomas D Alexander andS Santhakumari ldquoVariability heritability and correlation onstudies on the yield and quality characters of sugarcanerdquo IndianSugar vol 44 no 4 pp 243ndash249 1994
[42] J R Harlan J M de Wet and E G Price ldquoComparativeevolution of cerealsrdquo Evolution vol 27 no 2 pp 311ndash325 1973
[43] A Elings ldquoDurum wheat landraces from syria II patterns ofvariationrdquo Euphytica vol 54 no 3 pp 231ndash243 1991
[44] K Tato ldquoRainfall in Ethiopiardquo Ethiopian Geographical Journalvol 2 pp 28ndash36 1964
[45] A B L Stemler J R Harlan and J M J deWet ldquoThe sorghumsof ethiopiardquo Economic Botany vol 31 no 4 pp 446ndash460 1977
[46] P Annicchiarico L Pecetti and A B Damania ldquoRelationshipsbetween phenotypic variation and climatic factors at collectingsites in durum wheat landracesrdquo Hereditas vol 122 no 2 pp163ndash167 1995
[47] A B Damania L Pecetti C O Qualset and B O HumeidldquoDiversity and geographic distribution of adaptive traits inTriticum turgidum L (durum group) wheat landraces fromTurkeyrdquoGenetic Resources and Crop Evolution vol 43 no 5 pp409ndash422 1996
[48] A Ayana and E Bekele ldquoGeographical patterns of morpho-logical variation in sorghum (Sorghum bicolor (L) Moench)germplasm from Ethiopia and Eritrea quantitative charactersrdquoEuphytica vol 115 no 2 pp 91ndash104 2000
[49] D S Falconer Introduction to Quantitative Genetics JohnWilley and Sons Inc New York NY USA 3rd edition 1993
[50] R S Thorpe ldquoBiochemical analysis of geographical variationand racial affinitiesrdquo Biology Review vol 51 pp 407ndash452 1976
[51] M Farooq ldquoMorphological and agricultural characteristicsof sugarcane in relation to mechanized agriculture at NARCIslamabadrdquo Portu Rico Journal vol 1 pp 51ndash61 1989
[52] M Akhtar N N Elahi and M Ashraf ldquoEvaluation of exoticsugarcane varieties for agronomic characters and productivityrdquoPakistan Journal of Biological Sciences vol 4 no 1 pp 37ndash402001
[53] F R Piscitelli ldquoPrincipals caracteres exomorfologicos de cul-tures de cana de azucarrdquo TUC Iindustraly Agricos vol 7 pp 49ndash57 1994 (Spanish)
[54] A H D Brown and B S Weir ldquoMeasuring genetic variability inplant populationsrdquo in Isozyme in Plant Genetics Breeding S DTanksley and T J Orton Eds part A Elsevier Science 1983
[55] R Balakrishnan N V Nair and T V Sreenivasan ldquoA methodfor establishing a core collection of Saccharum officinarum Lgermplasm based on quantitative-morphological datardquo GeneticResources and Crop Evolution vol 47 no 1 pp 1ndash9 2000
[56] C A Nestor and B Q Reynaldo Sugarcane Variety Improve-ment in Southeast Asia and the Pacific for Enhanced andSustainable Productivity ndash Germplasm and Disease IndexingComponents Institute of Plant Breeding (IPB) College ofAgriculture (CA) UP Los Banos (UPLB) 2003
[57] A Maqbool J Muhammad and A Sagheer ldquoAgronomic traitsand morphological characteristics of some exotic varieties ofsugarcanerdquoPakistan Journal of Agricultural Research vol 19 no4 2006
suggests the advantage of a scheme of selection for morethan one character at a time The correlation between char-acters may arise from linkage or from developmental geneticinteractions with orwithout a purely phenotypic componentIt could arise also due to differential phenotypic plasticityof characters themselves Within the limit of experimentalerror and environmental effects high correlation coefficientsbetween characters may show that the characters sharesome common element of genetic control (ie pleiotropylinkage) between genes or else from independently controlledcharacters responding similarly to geographic variation inselection pressures [49] The interregion correlation coef-ficient between the characters measures the consistency oftheir patterns of regional variation while the intraregioncorrelation coefficient measures the association arising fromgenetic factors but is not affected by regional variation[50] Since this study showed significant positive correlationsintraregionally for some character combinations it wouldseem that common genetic control might be playing a role inbringing about correlations between the various charactersIt appeared that different response to regional variationwas playing a greater role than different genetic controlas shown by the many more significant and moderate tohigh correlation coefficients intraregionally than interregion-ally
322 Qualitative Characters The frequency distribution forthe 16 qualitative characters of germplasm samples by regionsin Ethiopia and of the exotic accessions is shown in Supple-mental Table 5 For the purpose of avoiding redundancy andimproving readability the discrete characters are presented bygrouping related traits together
BudCushion andRelativeDegree of BudExtension Evaluationof the presence and absence of bud cushion revealed thatgreat proportion (70) of sugarcane landraces from Ethiopiado not have bud cushion Majority of the sample germplasmin all regions belong to this phenotypic class Approximatelyone-half (53) of the collection have their bud extendedabove the growth ring followed by those (41) having budtouching the growth ring
Relative Bud Shape Among the phenotypic classes for thischaracter tall deltoid was dominant (31) followed by ovate(17) and narrow ovate (12) While local accessions inmost regions had tall deltoid bud shape 50 of the intro-duced accessions belong to two phenotypic classes roundwith central germpore (30) and 20 having pentagonalbud shape Equal proportions (19) of the samples fromBenshangul-Gumz were Ovate and Ovate with emarginatebasal wing
Canopy Structure Relative Plant Erectness Colour of theLeaves and Colour of the Exposed Rind Out of the eightphenotypic classes observed compact tip droopy was themost frequent (35) Open semidroopy and open droopyclasses had fairly equal distribution With regard to relativeplant erectness seventy-five percent of the germplasm werefound to fall in twophenotypic classes erect (41) and almost
erect (34) for the whole region and country Most of theaccessions (90) showed light green or green leaf colourbut 20 from introduced collections were greenish yellowAmong phenotypic classes for colour of the exposed rindbrownish yellow yellow brown and yellowish green were themost frequent with fairly equal distribution
Relative Shape of Dewlap Double crescent had larger fre-quency (22) while squarish descending and flaring typeof dewlap shape had fairly equal proportions Seventeenpercent of germplasm from Tigray region had squarishdeltoid dewlap shape
Relative Degree of Internode Alignment and Internode ShapeForty-four percent of the sample germplasm were slightlyzigzag while there was almost equal frequency for straightand zigzag internode alignment for the entire region andcountry Straight or nearly straight (slightly zigzag) internodealignment of cane stalks is a very important character formechanized farming and postharvest handling in sugarcane[51] Forty percent of the collections were observed havingconcave-convex type of internode shape while 29 wascylindrical and 14 conoidal Other classes had fairly equalproportion
Type of Auricle and Relative Shape of Ligule An equalproportion of the germplasm totally constituting 84 ofthe collection belong to four phenotypic classes namelythose with no auricle and others with transitional and shortand lanceolate auricles Sixteen percent of the collectionsalso showed equal frequency for the remaining classesBenshangul-Gumz and Tigray regions had each equal 13accessions with falcate type of auricle Sixty-seven percentof the collections belong to three ligule shape classes cres-cent with lozenge (27) broad-crescent (21) and linear-crescent (19) The germplasm had almost equal proportionfor the other phenotypic classes of this character Nineteenpercent of the accessions fromBenshangul-Gumz region haddeltoid shape of ligule
Stalk Corky Cracks Corky Patches Growth Cracks and BudGroove Eighty-nine percent of the germplasm across regionshad no stalk corky cracks with only eleven percent exhibitingthis character All the germplasm in Gambella Harari andSomali had no stalk corky cracks On the other hand eighty-one percent of the germplasm for the whole region showedthe presence of stalk corky patches All the local germplasmin Benshangul-Gumz Gambella and Tigray had stalk corkypatches whereas 75 of the germplasm from Harari Region67 from Somali Region and 24 from Oromia Regionhad no stalk corky patches In the entire region 75 ofthe germplasm had no stalk growth cracks with only 25showing this character Sixty-three percent of the germplasmwere found to have bud groove Howevermost of the samplesin Harari (75) Amhara (63) and Oromia (53) Regionshad no bud groove
The results of this study indicated the wide distributionof phenotypic classes for the characters considered whichindicates the existence of different races and combination
Advances in Agriculture 15
of races in local genotypes of different regions in EthiopiaThis diversity of germplasmbased onphenotypicmarkers canhelp sugarcane breeders in identifying the identity of geno-types as well as maintaining genetic diversity Differencesin morphological characters of different sugarcane varietieshave been reported [52] Piscitelli [53] showed importantexomorphological qualitative characters of sugarcane varietythat are not influenced by environmental factors and thus canbe used as a selection tool in any breeding program
The amount of phenotypic diversity estimates based onthe ShannonndashWeaver diversity index (H1015840) are shown inTable 9 The 16 characters differed in their distribution aswell as the amount of variation Individual traits showed adifferent pattern of variation among accessions Estimatesof H1015840 for individual characters varied from 049 for stalkcorky cracks to 100 for bud groove with an overall meanof 080 Most of the characters were highly polymorphicwhereas relative plant erectness and presence or absenceof stalk corky patches scored moderate values The highdiversity values for the characters showed a wide variabilityamong genotypes According to [54] this index is also usedin genetic resource studies as a convenient measure of bothallelic richness and allelic evenness when using geneticdata
The highest diversity computed was on presence orabsence of bud groove with a diversity index of 100indicating that this character is very variable in the localsugarcane germplasm On the other hand the descriptorcomputed with the lowest diversity index value (049) wasthe presence or absence of stalk corky cracks indicatingthat this character is selected in the development of thegenotypes In agreement with the present study [55] workingon S officinarum accessions from the world collection ofsugarcane germplasm obtained a low index of 025 and 035for presence or absence of stalk corky patches and cracksrespectively
Their report also showed high diversity index values forinternode alignment growth cracks internode shape budcushion bud extension bud groove and ligule shape Sim-ilarly [56] experimenting on field collections of sugarcaneaccessions in Philippine reported high ShannonndashWeaverdiversity for stalk growth cracks bud cushion stalk corkypatch stalk corky cracks bud extension and auricle outershape Unlike the present study they have reported high indexfor stalk corky cracksMedium index values were reported forstalk internode shape and low index values for ligule shapeand bud shape High variability of sugarcane varieties fordewlap shape was also reported in [57]
The H1015840 pooled across characters by region ranged from079 in Oromiya to 085 in Tigray with an overall aver-age of 081 (Table 8) Most of the regions showed highShannonndashWeaver diversity index The populations that hadthe highest H1015840 were that of Tigray and introduction The H1015840averaged over regions for different characters was found torange from 044 for presence or absence of stalk corky cracksto 094 for relative degree of internode alignment and budgroove with an overall average of 081 (Table 9)
Assessment of the ShannonndashWeaver index also showedsensitivity to both the number of descriptor classes and the
distribution within phenotypic classes in each region Amongregions the mean values for the ShannonndashWeaver index arenot significantly different but differences were found whenindividual characters were considered These observationssuggest that some regions have high diversity for particulartraits while in other regions selective pressures might havereduced the variation to certain genotypes
4 Conclusions
The large number of landraces observed during the cur-rent study demonstrates the existence of diverse geneticresources of sugarcane in Ethiopia The landraces wouldserve to broaden the genetic base of the available sugarcanegermplasm Generally distribution of the overall phenotypicdiversity among sugarcane landraces in Ethiopia is unevenOur results showed that there was a wide range of variationthat existed in the local sugarcane landraces studied both atregional and within region levels Selection and differenti-ation of types might have taken place along a geographicalpattern perhaps associated with climate and use The presentstudy contributes to updating sugarcane descriptors adoptedfrom USDA-ARS as well as Bioversity passport data forthe future collection and evaluation As Ethiopian SugarCorporation is in the course of launching breeding programthe observed substantial variation of landraces would enablesugarcane breeders to design and practice breeding andselection programs to improve the crop
Data Availability
The data used to support the findings of this study areavailable from the corresponding author upon request
Conflicts of Interest
The authors declare that they have no conflicts of interest
Acknowledgments
The authors are grateful for the financial grant of SugarCorporation of Ethiopia Many thanks are due to EthiopianBiodiversity Institute (EBI) for its technical advice duringcollection of the local sugarcane genotypes Sincere thanksare forwarded to Ethiopian smallholder farmers who pre-served sugarcane germplasm and shared their knowledgeand experience Regional Zonal and District (ldquoWeredardquo)Agriculture Bureaus across Ethiopia are acknowledged fortheir support during the study
Supplementary Materials
These supplementary materials contain different tables sup-porting the results of the study These include passport dataof sugarcane germplasm collected and related informationon the materials local sugarcane germplasm collected andused formorphological diversity study introduced sugarcane
16 Advances in Agriculture
Table9Sh
anno
n-WeaverD
iversityIndices(H1015840)a
ndmeanandsta
ndarderror(plusmnS
E)of
land
racesineach
region
basedon
16discretecharacters
Characters
Amha
raBe
nsha
ngul-G
umz
Introd
uced
Oromia
SNNP
Tigray
ELb
Total
Presence
orabsenceo
fbud
cushion(BUDCUSH
ION)
083
095
095
089
082
098
090
088
Relativ
edegreeo
fbud
extension(BUDEX
TEND)
077
079
093
074
072
087
080
078
Relativ
ebud
Shape(BU
DSH
APE
)078
096
089
081
085
078
084
078
Canop
ystructure(C
ANOPY
)089
098
095
081
078
096
089
084
Relativ
eshape
ofdewlap(D
EWLA
PSHAP)
087
094
092
090
082
096
090
086
Relativ
eplant
erectness(ER
ECT)
080
082
050
067
071
088
073
064
Relativ
edegreeo
finterno
dealignm
ent(IN
ALIGN)
096
076
098
097
098
100
094
098
Relativ
einterno
deshape(IN
SHAPE
)086
069
095
077
083
091
083
082
Color
oftheleaves(LE
AFC
OLO
R)087
071
090
064
074
071
076
076
Type
ofou
tera
uricle(A
URI
CLE
OUT)
080
092
087
079
083
093
086
082
Relativ
eshape
ofligule(LIGSH
APE
)086
094
082
085
090
083
087
083
Presence
orabsenceo
fstalkcorkyc
racks(ST
ALK
CORK
C)
044
054
035
055
051
025
044
049
Presence
orabsenceo
fstalkcorkyp
atches(STA
LKCORK
P)066
000
072
079
059
087
060
069
Presence
orabsenceo
fstalkgrow
thcracks
(STA
LKCRA
CK)
066
100
088
062
088
099
084
080
Color
ofthee
xposed
rind(R
INDCOLE
)085
096
089
084
086
076
086
085
Presence
orabsenceo
fbud
groo
ve(BUDGRO
OVE)
095
100
072
100
100
099
094
100
Means
080plusmn0
03
081plusmn0
06
083plusmn0
04
079plusmn0
03
080plusmn0
03
085plusmn0
05
081plusmn0
03
080
b EL=
averageo
fentire
locatio
n
Advances in Agriculture 17
varieties used for morphological diversity study and fre-quency distribution for 16 qualitative characters in sugarcaneby regions in Ethiopia and country of sources of introducedgenotypes (Supplementary Materials)
References
[1] ldquoSpreading the word about clean solutions from sugarcanrdquoAccessed 04072018 httpwwwsugarcaneorg
[2] Central Statistics Agency (CSA) ldquoAgricultural sample surveyArea and production of major crops private peasant holdingsrdquoStatistical Bulletin Addis Ababa Ethiopia 2017
[3] T Esayas Exploration and Collection Characterization GeneticDiversity Analysis and Association of Traits for Yield and YieldComponents of Sugarcane (Saccharumspp) in Ethiopia [MasterThesis] Haramaya University Dire Dawa Ethiopia 2014
[4] J R Harlan Crops and Man American Society of AgronomyCrop Science Society of America Madison WI USA 2ndedition 1992
[5] R W Allard Principles of Plant Breeding JohnWilley and SonsNew York NY USA 2nd edition 1999
[6] B T Roach and J Daniels ldquoA review of the origin and improve-ment of sugarcanerdquo in Proceedings of Coper sugar InternationalSugarcane Workshop pp 1ndash32 1987
[7] P P Patil S S Patil and D U Patel ldquoGenetic diversity in sug-arcane (Saccharum spp Complex)rdquo Journal of Pharmacognosyand Phytochemistry vol 6 no 6 pp 570ndash573 2017
[8] R B Singh B Singh andR K Singh ldquoStudy of genetic diversityof sugarcane (Saccharum) species and commercial varietiesthrough TRAP molecular markersrdquo Indian Journal of PlantPhysiology vol 22 no 3 pp 332ndash338 2017
[9] R W Allard ldquoPopulation structure and sampling methodsrdquo inGenetic Resources in PlantsmdashTheir Exploration and Conserva-tion O H Frankel and E Bennett Eds IBP Handbook no 11pp 97ndash107 FA Davis Company Philadelphia PA USA 1970
[10] D R Marshall and A H D Brown ldquoOptimum samplingstrategies in genetic conservationrdquo inCrop Genetic Resources forToday and Tomorrow O H Frankel and J G Hawkes Eds pp53ndash80 Cambridge University Press 1975
[11] J G Hawkes Crop Genetic Resources Field Collection ManualPudoc Wagnningen Netherland 1980
[12] J G Hawkes The Diversity of Crop Plants Harvard UniversityPress London England 1983
[13] B P K Yerima Manual for Good Laboratory Practice Foodand Agriculture Organization of United Nation Addis AbabaEthiopia 1993
[14] A Walkley and I A Black ldquoAn examination of the degtjareffmethod for determining soil organic matter and a proposedmodification of the chromic acid titrationmethodrdquo Soil Sciencevol 37 no 1 pp 29ndash38 1934
[15] S R Olsen and L E Sommers ldquoPhosphorusrdquo in AgronomyMonograph A L Page et al Ed Methods of soil analysis part2 pp 403ndash430 ASA and SSSAMadisonWI USA 2nd edition1982
[16] R H Bray and L T Kurtz ldquoDetermination of total organic andavailable forms of phosphorus in soilsrdquo Soil Science vol 59 no1 pp 39ndash46 1945
[17] S Sahlemedhin andB Taye ldquoProcedures for soil and plant anal-ysisrdquo Tech Rep 74 National Soil Research Center EthiopianAgricultural Research Organization 2000
[18] F C Stickler S Weaden and AW Pauli ldquoLeaf area determina-tion in grain sorghumrdquo Journal of Agronomy and Crop Sciencevol 53 pp 187-188 1961
[19] Munsell Color ChartThe Munsell Book of Color Matte Collec-tion Munsell Color Chart NewWindsor NY USA 1990
[20] H O Hartley ldquoThe maximum F-ratio as a short cut test forheterogeneity of variancesrdquo Biometrika vol 37 pp 308ndash3121950
[21] L Pecetti P Annicchiarico and A B Damania ldquoBiodiversity ina germplasm collection of durumwheatrdquo Euphytica vol 60 no3 pp 229ndash238 1992
[22] L Pecetti and A B Damania ldquoGeographic variation intetraploid wheat (Triticum turgidum ssp turgidum convardurum) landraces from two provinces in Ethiopiardquo GeneticResources and Crop Evolution vol 43 no 5 pp 395ndash407 1996
[23] S K Jain C O Qualset G M Bhatt and K K Wu ldquoGeo-graphical patterns of phenotypic diversity in a world collectionof durumwheatsrdquoCrop Science vol 15 no 5 pp 700ndash704 1975
[24] K K Nkongolo L Chinthu M Malusi and Z Vokhiwa ldquoPar-ticipatory variety selection and characterization of sorghum(Sorghum bicolour (L) Moench) elite accessions from Malaw-ian gene pool using farmer and breeder knowledgerdquo AfricanJournal of Agricultural Research vol 3 pp 273ndash283 2008
[25] M B Semira ldquoInnovative methods and metrics for agricul-ture and nutrition actions soil quality and human nutritionin Ethiopiardquo Leverhulme Center for Integrative Research onAgriculture and Health (LCIRAH) 2017
[26] I A Khan A Khatri M A Siddiqui G S Nizamani and SRaza ldquoPerformance of promising sugarcane clone for yield andquality traits in different ecological zones of Sindhirdquo PakistanJournal of Botany vol 36 no 1 pp 83ndash92 2004
[27] M F Perera M E Arias D Costilla et al ldquoGenetic diversityassessment and genotype identification in sugarcane based onDNAmarkers andmorphological traitsrdquo Euphytica vol 185 no3 pp 491ndash510 2012
[28] C G Neil M Kay and C C Jack ldquoDiversity among mainlandUSA sugarcane cultivars examined by SSR genotypingrdquo Journalof the American Society of Sugar Cane Technologists vol 29 pp36ndash52 2009
[29] S SchenckMWCrepeauK KWu P HMooreQ Yu andRMing ldquoGenetic diversity and relationships in native HawaiianSaccharum officinarum sugarcanerdquo Journal of Heredity vol 95no 4 pp 327ndash331 2004
[30] R Gebretsadik H Shimelis M D Laing P Tongoona andN Mandefro ldquoA diagnostic appraisal of the sorghum farmingsystem and breeding priorities in Striga infested agro-ecologiesof Ethiopiardquo Agricultural Systems vol 123 pp 54ndash61 2014
[31] L E V Cesar A T Bruzi J A R Nunes et al ldquoGeneticdivergence of sugarcane varieties collected in the region ofLavras Minas Gerais Brazilrdquo Genetics and Molecular Researchvol 14 no 4 pp 14043ndash14049 2015
[32] C Grenier P J Bramel J A Dahlberg et al ldquoSorghums of theSudan analysis of regional diversity and distributionrdquo GeneticResources and Crop Evolution vol 51 no 5 pp 489ndash500 2004
[33] N V Nair R Balakrishnan and T V Sreenivasan ldquoVariabilityfor quantitative traits in exotic hybrid germplasm of sugarcanerdquoGenetic Resources and Crop Evolution vol 45 no 5 pp 459ndash464 1998
[34] R Nagarajan S Alarmelu and R M Shanthi ldquoStudies onvariation in interspecific hybrids of Saccharumrdquo Sugar Tech vol2 no 3 pp 42ndash46 2000
18 Advances in Agriculture
[35] D N Kamat and J R P Singh ldquoVariability in Sugarcane underrain fed conditionrdquo Sugar Technology Short Communicationsvol 3 no 1-2 pp 65ndash67 2001
[36] B S Rishipal A S M Chaudhary and S P Kadian ldquoStudieson character interrelationship in clonal generation of sugarcane(Saccharum complex hybrids)rdquo Indian Sugar vol 47 no 11 pp907ndash911 1998
[37] S Thippeswamy S T Kajjidoni P M Salimath and J V GoudldquoCorrelation and path analysis for cane yield juice quality andtheir component traits in sugarcanerdquo Short Communication vol5 no 1-2 pp 65ndash72 2003
[38] P S Verma S Pal and N K Karma ldquoGenetic variability andcorrelation studies in sugarcanerdquo Indian Sugar vol 49 pp 125ndash128 1999
[39] H A Esechie J W Maranville and W M Ross ldquoRelationshipof stalk morphology and chemical composition to lodgingresistance in sorghumrdquo Crop Science vol 17 no 4 p 609 1977
[40] P K Das B C Jena and N Nayak ldquoCharacter association andpath analysis of sugar yield in sugarcanerdquo Indian Sugar vol 46no 10 pp 805ndash808 1997
[41] K J Sreekumar M Kuriakose M Thomas D Alexander andS Santhakumari ldquoVariability heritability and correlation onstudies on the yield and quality characters of sugarcanerdquo IndianSugar vol 44 no 4 pp 243ndash249 1994
[42] J R Harlan J M de Wet and E G Price ldquoComparativeevolution of cerealsrdquo Evolution vol 27 no 2 pp 311ndash325 1973
[43] A Elings ldquoDurum wheat landraces from syria II patterns ofvariationrdquo Euphytica vol 54 no 3 pp 231ndash243 1991
[44] K Tato ldquoRainfall in Ethiopiardquo Ethiopian Geographical Journalvol 2 pp 28ndash36 1964
[45] A B L Stemler J R Harlan and J M J deWet ldquoThe sorghumsof ethiopiardquo Economic Botany vol 31 no 4 pp 446ndash460 1977
[46] P Annicchiarico L Pecetti and A B Damania ldquoRelationshipsbetween phenotypic variation and climatic factors at collectingsites in durum wheat landracesrdquo Hereditas vol 122 no 2 pp163ndash167 1995
[47] A B Damania L Pecetti C O Qualset and B O HumeidldquoDiversity and geographic distribution of adaptive traits inTriticum turgidum L (durum group) wheat landraces fromTurkeyrdquoGenetic Resources and Crop Evolution vol 43 no 5 pp409ndash422 1996
[48] A Ayana and E Bekele ldquoGeographical patterns of morpho-logical variation in sorghum (Sorghum bicolor (L) Moench)germplasm from Ethiopia and Eritrea quantitative charactersrdquoEuphytica vol 115 no 2 pp 91ndash104 2000
[49] D S Falconer Introduction to Quantitative Genetics JohnWilley and Sons Inc New York NY USA 3rd edition 1993
[50] R S Thorpe ldquoBiochemical analysis of geographical variationand racial affinitiesrdquo Biology Review vol 51 pp 407ndash452 1976
[51] M Farooq ldquoMorphological and agricultural characteristicsof sugarcane in relation to mechanized agriculture at NARCIslamabadrdquo Portu Rico Journal vol 1 pp 51ndash61 1989
[52] M Akhtar N N Elahi and M Ashraf ldquoEvaluation of exoticsugarcane varieties for agronomic characters and productivityrdquoPakistan Journal of Biological Sciences vol 4 no 1 pp 37ndash402001
[53] F R Piscitelli ldquoPrincipals caracteres exomorfologicos de cul-tures de cana de azucarrdquo TUC Iindustraly Agricos vol 7 pp 49ndash57 1994 (Spanish)
[54] A H D Brown and B S Weir ldquoMeasuring genetic variability inplant populationsrdquo in Isozyme in Plant Genetics Breeding S DTanksley and T J Orton Eds part A Elsevier Science 1983
[55] R Balakrishnan N V Nair and T V Sreenivasan ldquoA methodfor establishing a core collection of Saccharum officinarum Lgermplasm based on quantitative-morphological datardquo GeneticResources and Crop Evolution vol 47 no 1 pp 1ndash9 2000
[56] C A Nestor and B Q Reynaldo Sugarcane Variety Improve-ment in Southeast Asia and the Pacific for Enhanced andSustainable Productivity ndash Germplasm and Disease IndexingComponents Institute of Plant Breeding (IPB) College ofAgriculture (CA) UP Los Banos (UPLB) 2003
[57] A Maqbool J Muhammad and A Sagheer ldquoAgronomic traitsand morphological characteristics of some exotic varieties ofsugarcanerdquoPakistan Journal of Agricultural Research vol 19 no4 2006
suggests the advantage of a scheme of selection for morethan one character at a time The correlation between char-acters may arise from linkage or from developmental geneticinteractions with orwithout a purely phenotypic componentIt could arise also due to differential phenotypic plasticityof characters themselves Within the limit of experimentalerror and environmental effects high correlation coefficientsbetween characters may show that the characters sharesome common element of genetic control (ie pleiotropylinkage) between genes or else from independently controlledcharacters responding similarly to geographic variation inselection pressures [49] The interregion correlation coef-ficient between the characters measures the consistency oftheir patterns of regional variation while the intraregioncorrelation coefficient measures the association arising fromgenetic factors but is not affected by regional variation[50] Since this study showed significant positive correlationsintraregionally for some character combinations it wouldseem that common genetic control might be playing a role inbringing about correlations between the various charactersIt appeared that different response to regional variationwas playing a greater role than different genetic controlas shown by the many more significant and moderate tohigh correlation coefficients intraregionally than interregion-ally
322 Qualitative Characters The frequency distribution forthe 16 qualitative characters of germplasm samples by regionsin Ethiopia and of the exotic accessions is shown in Supple-mental Table 5 For the purpose of avoiding redundancy andimproving readability the discrete characters are presented bygrouping related traits together
BudCushion andRelativeDegree of BudExtension Evaluationof the presence and absence of bud cushion revealed thatgreat proportion (70) of sugarcane landraces from Ethiopiado not have bud cushion Majority of the sample germplasmin all regions belong to this phenotypic class Approximatelyone-half (53) of the collection have their bud extendedabove the growth ring followed by those (41) having budtouching the growth ring
Relative Bud Shape Among the phenotypic classes for thischaracter tall deltoid was dominant (31) followed by ovate(17) and narrow ovate (12) While local accessions inmost regions had tall deltoid bud shape 50 of the intro-duced accessions belong to two phenotypic classes roundwith central germpore (30) and 20 having pentagonalbud shape Equal proportions (19) of the samples fromBenshangul-Gumz were Ovate and Ovate with emarginatebasal wing
Canopy Structure Relative Plant Erectness Colour of theLeaves and Colour of the Exposed Rind Out of the eightphenotypic classes observed compact tip droopy was themost frequent (35) Open semidroopy and open droopyclasses had fairly equal distribution With regard to relativeplant erectness seventy-five percent of the germplasm werefound to fall in twophenotypic classes erect (41) and almost
erect (34) for the whole region and country Most of theaccessions (90) showed light green or green leaf colourbut 20 from introduced collections were greenish yellowAmong phenotypic classes for colour of the exposed rindbrownish yellow yellow brown and yellowish green were themost frequent with fairly equal distribution
Relative Shape of Dewlap Double crescent had larger fre-quency (22) while squarish descending and flaring typeof dewlap shape had fairly equal proportions Seventeenpercent of germplasm from Tigray region had squarishdeltoid dewlap shape
Relative Degree of Internode Alignment and Internode ShapeForty-four percent of the sample germplasm were slightlyzigzag while there was almost equal frequency for straightand zigzag internode alignment for the entire region andcountry Straight or nearly straight (slightly zigzag) internodealignment of cane stalks is a very important character formechanized farming and postharvest handling in sugarcane[51] Forty percent of the collections were observed havingconcave-convex type of internode shape while 29 wascylindrical and 14 conoidal Other classes had fairly equalproportion
Type of Auricle and Relative Shape of Ligule An equalproportion of the germplasm totally constituting 84 ofthe collection belong to four phenotypic classes namelythose with no auricle and others with transitional and shortand lanceolate auricles Sixteen percent of the collectionsalso showed equal frequency for the remaining classesBenshangul-Gumz and Tigray regions had each equal 13accessions with falcate type of auricle Sixty-seven percentof the collections belong to three ligule shape classes cres-cent with lozenge (27) broad-crescent (21) and linear-crescent (19) The germplasm had almost equal proportionfor the other phenotypic classes of this character Nineteenpercent of the accessions fromBenshangul-Gumz region haddeltoid shape of ligule
Stalk Corky Cracks Corky Patches Growth Cracks and BudGroove Eighty-nine percent of the germplasm across regionshad no stalk corky cracks with only eleven percent exhibitingthis character All the germplasm in Gambella Harari andSomali had no stalk corky cracks On the other hand eighty-one percent of the germplasm for the whole region showedthe presence of stalk corky patches All the local germplasmin Benshangul-Gumz Gambella and Tigray had stalk corkypatches whereas 75 of the germplasm from Harari Region67 from Somali Region and 24 from Oromia Regionhad no stalk corky patches In the entire region 75 ofthe germplasm had no stalk growth cracks with only 25showing this character Sixty-three percent of the germplasmwere found to have bud groove Howevermost of the samplesin Harari (75) Amhara (63) and Oromia (53) Regionshad no bud groove
The results of this study indicated the wide distributionof phenotypic classes for the characters considered whichindicates the existence of different races and combination
Advances in Agriculture 15
of races in local genotypes of different regions in EthiopiaThis diversity of germplasmbased onphenotypicmarkers canhelp sugarcane breeders in identifying the identity of geno-types as well as maintaining genetic diversity Differencesin morphological characters of different sugarcane varietieshave been reported [52] Piscitelli [53] showed importantexomorphological qualitative characters of sugarcane varietythat are not influenced by environmental factors and thus canbe used as a selection tool in any breeding program
The amount of phenotypic diversity estimates based onthe ShannonndashWeaver diversity index (H1015840) are shown inTable 9 The 16 characters differed in their distribution aswell as the amount of variation Individual traits showed adifferent pattern of variation among accessions Estimatesof H1015840 for individual characters varied from 049 for stalkcorky cracks to 100 for bud groove with an overall meanof 080 Most of the characters were highly polymorphicwhereas relative plant erectness and presence or absenceof stalk corky patches scored moderate values The highdiversity values for the characters showed a wide variabilityamong genotypes According to [54] this index is also usedin genetic resource studies as a convenient measure of bothallelic richness and allelic evenness when using geneticdata
The highest diversity computed was on presence orabsence of bud groove with a diversity index of 100indicating that this character is very variable in the localsugarcane germplasm On the other hand the descriptorcomputed with the lowest diversity index value (049) wasthe presence or absence of stalk corky cracks indicatingthat this character is selected in the development of thegenotypes In agreement with the present study [55] workingon S officinarum accessions from the world collection ofsugarcane germplasm obtained a low index of 025 and 035for presence or absence of stalk corky patches and cracksrespectively
Their report also showed high diversity index values forinternode alignment growth cracks internode shape budcushion bud extension bud groove and ligule shape Sim-ilarly [56] experimenting on field collections of sugarcaneaccessions in Philippine reported high ShannonndashWeaverdiversity for stalk growth cracks bud cushion stalk corkypatch stalk corky cracks bud extension and auricle outershape Unlike the present study they have reported high indexfor stalk corky cracksMedium index values were reported forstalk internode shape and low index values for ligule shapeand bud shape High variability of sugarcane varieties fordewlap shape was also reported in [57]
The H1015840 pooled across characters by region ranged from079 in Oromiya to 085 in Tigray with an overall aver-age of 081 (Table 8) Most of the regions showed highShannonndashWeaver diversity index The populations that hadthe highest H1015840 were that of Tigray and introduction The H1015840averaged over regions for different characters was found torange from 044 for presence or absence of stalk corky cracksto 094 for relative degree of internode alignment and budgroove with an overall average of 081 (Table 9)
Assessment of the ShannonndashWeaver index also showedsensitivity to both the number of descriptor classes and the
distribution within phenotypic classes in each region Amongregions the mean values for the ShannonndashWeaver index arenot significantly different but differences were found whenindividual characters were considered These observationssuggest that some regions have high diversity for particulartraits while in other regions selective pressures might havereduced the variation to certain genotypes
4 Conclusions
The large number of landraces observed during the cur-rent study demonstrates the existence of diverse geneticresources of sugarcane in Ethiopia The landraces wouldserve to broaden the genetic base of the available sugarcanegermplasm Generally distribution of the overall phenotypicdiversity among sugarcane landraces in Ethiopia is unevenOur results showed that there was a wide range of variationthat existed in the local sugarcane landraces studied both atregional and within region levels Selection and differenti-ation of types might have taken place along a geographicalpattern perhaps associated with climate and use The presentstudy contributes to updating sugarcane descriptors adoptedfrom USDA-ARS as well as Bioversity passport data forthe future collection and evaluation As Ethiopian SugarCorporation is in the course of launching breeding programthe observed substantial variation of landraces would enablesugarcane breeders to design and practice breeding andselection programs to improve the crop
Data Availability
The data used to support the findings of this study areavailable from the corresponding author upon request
Conflicts of Interest
The authors declare that they have no conflicts of interest
Acknowledgments
The authors are grateful for the financial grant of SugarCorporation of Ethiopia Many thanks are due to EthiopianBiodiversity Institute (EBI) for its technical advice duringcollection of the local sugarcane genotypes Sincere thanksare forwarded to Ethiopian smallholder farmers who pre-served sugarcane germplasm and shared their knowledgeand experience Regional Zonal and District (ldquoWeredardquo)Agriculture Bureaus across Ethiopia are acknowledged fortheir support during the study
Supplementary Materials
These supplementary materials contain different tables sup-porting the results of the study These include passport dataof sugarcane germplasm collected and related informationon the materials local sugarcane germplasm collected andused formorphological diversity study introduced sugarcane
16 Advances in Agriculture
Table9Sh
anno
n-WeaverD
iversityIndices(H1015840)a
ndmeanandsta
ndarderror(plusmnS
E)of
land
racesineach
region
basedon
16discretecharacters
Characters
Amha
raBe
nsha
ngul-G
umz
Introd
uced
Oromia
SNNP
Tigray
ELb
Total
Presence
orabsenceo
fbud
cushion(BUDCUSH
ION)
083
095
095
089
082
098
090
088
Relativ
edegreeo
fbud
extension(BUDEX
TEND)
077
079
093
074
072
087
080
078
Relativ
ebud
Shape(BU
DSH
APE
)078
096
089
081
085
078
084
078
Canop
ystructure(C
ANOPY
)089
098
095
081
078
096
089
084
Relativ
eshape
ofdewlap(D
EWLA
PSHAP)
087
094
092
090
082
096
090
086
Relativ
eplant
erectness(ER
ECT)
080
082
050
067
071
088
073
064
Relativ
edegreeo
finterno
dealignm
ent(IN
ALIGN)
096
076
098
097
098
100
094
098
Relativ
einterno
deshape(IN
SHAPE
)086
069
095
077
083
091
083
082
Color
oftheleaves(LE
AFC
OLO
R)087
071
090
064
074
071
076
076
Type
ofou
tera
uricle(A
URI
CLE
OUT)
080
092
087
079
083
093
086
082
Relativ
eshape
ofligule(LIGSH
APE
)086
094
082
085
090
083
087
083
Presence
orabsenceo
fstalkcorkyc
racks(ST
ALK
CORK
C)
044
054
035
055
051
025
044
049
Presence
orabsenceo
fstalkcorkyp
atches(STA
LKCORK
P)066
000
072
079
059
087
060
069
Presence
orabsenceo
fstalkgrow
thcracks
(STA
LKCRA
CK)
066
100
088
062
088
099
084
080
Color
ofthee
xposed
rind(R
INDCOLE
)085
096
089
084
086
076
086
085
Presence
orabsenceo
fbud
groo
ve(BUDGRO
OVE)
095
100
072
100
100
099
094
100
Means
080plusmn0
03
081plusmn0
06
083plusmn0
04
079plusmn0
03
080plusmn0
03
085plusmn0
05
081plusmn0
03
080
b EL=
averageo
fentire
locatio
n
Advances in Agriculture 17
varieties used for morphological diversity study and fre-quency distribution for 16 qualitative characters in sugarcaneby regions in Ethiopia and country of sources of introducedgenotypes (Supplementary Materials)
References
[1] ldquoSpreading the word about clean solutions from sugarcanrdquoAccessed 04072018 httpwwwsugarcaneorg
[2] Central Statistics Agency (CSA) ldquoAgricultural sample surveyArea and production of major crops private peasant holdingsrdquoStatistical Bulletin Addis Ababa Ethiopia 2017
[3] T Esayas Exploration and Collection Characterization GeneticDiversity Analysis and Association of Traits for Yield and YieldComponents of Sugarcane (Saccharumspp) in Ethiopia [MasterThesis] Haramaya University Dire Dawa Ethiopia 2014
[4] J R Harlan Crops and Man American Society of AgronomyCrop Science Society of America Madison WI USA 2ndedition 1992
[5] R W Allard Principles of Plant Breeding JohnWilley and SonsNew York NY USA 2nd edition 1999
[6] B T Roach and J Daniels ldquoA review of the origin and improve-ment of sugarcanerdquo in Proceedings of Coper sugar InternationalSugarcane Workshop pp 1ndash32 1987
[7] P P Patil S S Patil and D U Patel ldquoGenetic diversity in sug-arcane (Saccharum spp Complex)rdquo Journal of Pharmacognosyand Phytochemistry vol 6 no 6 pp 570ndash573 2017
[8] R B Singh B Singh andR K Singh ldquoStudy of genetic diversityof sugarcane (Saccharum) species and commercial varietiesthrough TRAP molecular markersrdquo Indian Journal of PlantPhysiology vol 22 no 3 pp 332ndash338 2017
[9] R W Allard ldquoPopulation structure and sampling methodsrdquo inGenetic Resources in PlantsmdashTheir Exploration and Conserva-tion O H Frankel and E Bennett Eds IBP Handbook no 11pp 97ndash107 FA Davis Company Philadelphia PA USA 1970
[10] D R Marshall and A H D Brown ldquoOptimum samplingstrategies in genetic conservationrdquo inCrop Genetic Resources forToday and Tomorrow O H Frankel and J G Hawkes Eds pp53ndash80 Cambridge University Press 1975
[11] J G Hawkes Crop Genetic Resources Field Collection ManualPudoc Wagnningen Netherland 1980
[12] J G Hawkes The Diversity of Crop Plants Harvard UniversityPress London England 1983
[13] B P K Yerima Manual for Good Laboratory Practice Foodand Agriculture Organization of United Nation Addis AbabaEthiopia 1993
[14] A Walkley and I A Black ldquoAn examination of the degtjareffmethod for determining soil organic matter and a proposedmodification of the chromic acid titrationmethodrdquo Soil Sciencevol 37 no 1 pp 29ndash38 1934
[15] S R Olsen and L E Sommers ldquoPhosphorusrdquo in AgronomyMonograph A L Page et al Ed Methods of soil analysis part2 pp 403ndash430 ASA and SSSAMadisonWI USA 2nd edition1982
[16] R H Bray and L T Kurtz ldquoDetermination of total organic andavailable forms of phosphorus in soilsrdquo Soil Science vol 59 no1 pp 39ndash46 1945
[17] S Sahlemedhin andB Taye ldquoProcedures for soil and plant anal-ysisrdquo Tech Rep 74 National Soil Research Center EthiopianAgricultural Research Organization 2000
[18] F C Stickler S Weaden and AW Pauli ldquoLeaf area determina-tion in grain sorghumrdquo Journal of Agronomy and Crop Sciencevol 53 pp 187-188 1961
[19] Munsell Color ChartThe Munsell Book of Color Matte Collec-tion Munsell Color Chart NewWindsor NY USA 1990
[20] H O Hartley ldquoThe maximum F-ratio as a short cut test forheterogeneity of variancesrdquo Biometrika vol 37 pp 308ndash3121950
[21] L Pecetti P Annicchiarico and A B Damania ldquoBiodiversity ina germplasm collection of durumwheatrdquo Euphytica vol 60 no3 pp 229ndash238 1992
[22] L Pecetti and A B Damania ldquoGeographic variation intetraploid wheat (Triticum turgidum ssp turgidum convardurum) landraces from two provinces in Ethiopiardquo GeneticResources and Crop Evolution vol 43 no 5 pp 395ndash407 1996
[23] S K Jain C O Qualset G M Bhatt and K K Wu ldquoGeo-graphical patterns of phenotypic diversity in a world collectionof durumwheatsrdquoCrop Science vol 15 no 5 pp 700ndash704 1975
[24] K K Nkongolo L Chinthu M Malusi and Z Vokhiwa ldquoPar-ticipatory variety selection and characterization of sorghum(Sorghum bicolour (L) Moench) elite accessions from Malaw-ian gene pool using farmer and breeder knowledgerdquo AfricanJournal of Agricultural Research vol 3 pp 273ndash283 2008
[25] M B Semira ldquoInnovative methods and metrics for agricul-ture and nutrition actions soil quality and human nutritionin Ethiopiardquo Leverhulme Center for Integrative Research onAgriculture and Health (LCIRAH) 2017
[26] I A Khan A Khatri M A Siddiqui G S Nizamani and SRaza ldquoPerformance of promising sugarcane clone for yield andquality traits in different ecological zones of Sindhirdquo PakistanJournal of Botany vol 36 no 1 pp 83ndash92 2004
[27] M F Perera M E Arias D Costilla et al ldquoGenetic diversityassessment and genotype identification in sugarcane based onDNAmarkers andmorphological traitsrdquo Euphytica vol 185 no3 pp 491ndash510 2012
[28] C G Neil M Kay and C C Jack ldquoDiversity among mainlandUSA sugarcane cultivars examined by SSR genotypingrdquo Journalof the American Society of Sugar Cane Technologists vol 29 pp36ndash52 2009
[29] S SchenckMWCrepeauK KWu P HMooreQ Yu andRMing ldquoGenetic diversity and relationships in native HawaiianSaccharum officinarum sugarcanerdquo Journal of Heredity vol 95no 4 pp 327ndash331 2004
[30] R Gebretsadik H Shimelis M D Laing P Tongoona andN Mandefro ldquoA diagnostic appraisal of the sorghum farmingsystem and breeding priorities in Striga infested agro-ecologiesof Ethiopiardquo Agricultural Systems vol 123 pp 54ndash61 2014
[31] L E V Cesar A T Bruzi J A R Nunes et al ldquoGeneticdivergence of sugarcane varieties collected in the region ofLavras Minas Gerais Brazilrdquo Genetics and Molecular Researchvol 14 no 4 pp 14043ndash14049 2015
[32] C Grenier P J Bramel J A Dahlberg et al ldquoSorghums of theSudan analysis of regional diversity and distributionrdquo GeneticResources and Crop Evolution vol 51 no 5 pp 489ndash500 2004
[33] N V Nair R Balakrishnan and T V Sreenivasan ldquoVariabilityfor quantitative traits in exotic hybrid germplasm of sugarcanerdquoGenetic Resources and Crop Evolution vol 45 no 5 pp 459ndash464 1998
[34] R Nagarajan S Alarmelu and R M Shanthi ldquoStudies onvariation in interspecific hybrids of Saccharumrdquo Sugar Tech vol2 no 3 pp 42ndash46 2000
18 Advances in Agriculture
[35] D N Kamat and J R P Singh ldquoVariability in Sugarcane underrain fed conditionrdquo Sugar Technology Short Communicationsvol 3 no 1-2 pp 65ndash67 2001
[36] B S Rishipal A S M Chaudhary and S P Kadian ldquoStudieson character interrelationship in clonal generation of sugarcane(Saccharum complex hybrids)rdquo Indian Sugar vol 47 no 11 pp907ndash911 1998
[37] S Thippeswamy S T Kajjidoni P M Salimath and J V GoudldquoCorrelation and path analysis for cane yield juice quality andtheir component traits in sugarcanerdquo Short Communication vol5 no 1-2 pp 65ndash72 2003
[38] P S Verma S Pal and N K Karma ldquoGenetic variability andcorrelation studies in sugarcanerdquo Indian Sugar vol 49 pp 125ndash128 1999
[39] H A Esechie J W Maranville and W M Ross ldquoRelationshipof stalk morphology and chemical composition to lodgingresistance in sorghumrdquo Crop Science vol 17 no 4 p 609 1977
[40] P K Das B C Jena and N Nayak ldquoCharacter association andpath analysis of sugar yield in sugarcanerdquo Indian Sugar vol 46no 10 pp 805ndash808 1997
[41] K J Sreekumar M Kuriakose M Thomas D Alexander andS Santhakumari ldquoVariability heritability and correlation onstudies on the yield and quality characters of sugarcanerdquo IndianSugar vol 44 no 4 pp 243ndash249 1994
[42] J R Harlan J M de Wet and E G Price ldquoComparativeevolution of cerealsrdquo Evolution vol 27 no 2 pp 311ndash325 1973
[43] A Elings ldquoDurum wheat landraces from syria II patterns ofvariationrdquo Euphytica vol 54 no 3 pp 231ndash243 1991
[44] K Tato ldquoRainfall in Ethiopiardquo Ethiopian Geographical Journalvol 2 pp 28ndash36 1964
[45] A B L Stemler J R Harlan and J M J deWet ldquoThe sorghumsof ethiopiardquo Economic Botany vol 31 no 4 pp 446ndash460 1977
[46] P Annicchiarico L Pecetti and A B Damania ldquoRelationshipsbetween phenotypic variation and climatic factors at collectingsites in durum wheat landracesrdquo Hereditas vol 122 no 2 pp163ndash167 1995
[47] A B Damania L Pecetti C O Qualset and B O HumeidldquoDiversity and geographic distribution of adaptive traits inTriticum turgidum L (durum group) wheat landraces fromTurkeyrdquoGenetic Resources and Crop Evolution vol 43 no 5 pp409ndash422 1996
[48] A Ayana and E Bekele ldquoGeographical patterns of morpho-logical variation in sorghum (Sorghum bicolor (L) Moench)germplasm from Ethiopia and Eritrea quantitative charactersrdquoEuphytica vol 115 no 2 pp 91ndash104 2000
[49] D S Falconer Introduction to Quantitative Genetics JohnWilley and Sons Inc New York NY USA 3rd edition 1993
[50] R S Thorpe ldquoBiochemical analysis of geographical variationand racial affinitiesrdquo Biology Review vol 51 pp 407ndash452 1976
[51] M Farooq ldquoMorphological and agricultural characteristicsof sugarcane in relation to mechanized agriculture at NARCIslamabadrdquo Portu Rico Journal vol 1 pp 51ndash61 1989
[52] M Akhtar N N Elahi and M Ashraf ldquoEvaluation of exoticsugarcane varieties for agronomic characters and productivityrdquoPakistan Journal of Biological Sciences vol 4 no 1 pp 37ndash402001
[53] F R Piscitelli ldquoPrincipals caracteres exomorfologicos de cul-tures de cana de azucarrdquo TUC Iindustraly Agricos vol 7 pp 49ndash57 1994 (Spanish)
[54] A H D Brown and B S Weir ldquoMeasuring genetic variability inplant populationsrdquo in Isozyme in Plant Genetics Breeding S DTanksley and T J Orton Eds part A Elsevier Science 1983
[55] R Balakrishnan N V Nair and T V Sreenivasan ldquoA methodfor establishing a core collection of Saccharum officinarum Lgermplasm based on quantitative-morphological datardquo GeneticResources and Crop Evolution vol 47 no 1 pp 1ndash9 2000
[56] C A Nestor and B Q Reynaldo Sugarcane Variety Improve-ment in Southeast Asia and the Pacific for Enhanced andSustainable Productivity ndash Germplasm and Disease IndexingComponents Institute of Plant Breeding (IPB) College ofAgriculture (CA) UP Los Banos (UPLB) 2003
[57] A Maqbool J Muhammad and A Sagheer ldquoAgronomic traitsand morphological characteristics of some exotic varieties ofsugarcanerdquoPakistan Journal of Agricultural Research vol 19 no4 2006
suggests the advantage of a scheme of selection for morethan one character at a time The correlation between char-acters may arise from linkage or from developmental geneticinteractions with orwithout a purely phenotypic componentIt could arise also due to differential phenotypic plasticityof characters themselves Within the limit of experimentalerror and environmental effects high correlation coefficientsbetween characters may show that the characters sharesome common element of genetic control (ie pleiotropylinkage) between genes or else from independently controlledcharacters responding similarly to geographic variation inselection pressures [49] The interregion correlation coef-ficient between the characters measures the consistency oftheir patterns of regional variation while the intraregioncorrelation coefficient measures the association arising fromgenetic factors but is not affected by regional variation[50] Since this study showed significant positive correlationsintraregionally for some character combinations it wouldseem that common genetic control might be playing a role inbringing about correlations between the various charactersIt appeared that different response to regional variationwas playing a greater role than different genetic controlas shown by the many more significant and moderate tohigh correlation coefficients intraregionally than interregion-ally
322 Qualitative Characters The frequency distribution forthe 16 qualitative characters of germplasm samples by regionsin Ethiopia and of the exotic accessions is shown in Supple-mental Table 5 For the purpose of avoiding redundancy andimproving readability the discrete characters are presented bygrouping related traits together
BudCushion andRelativeDegree of BudExtension Evaluationof the presence and absence of bud cushion revealed thatgreat proportion (70) of sugarcane landraces from Ethiopiado not have bud cushion Majority of the sample germplasmin all regions belong to this phenotypic class Approximatelyone-half (53) of the collection have their bud extendedabove the growth ring followed by those (41) having budtouching the growth ring
Relative Bud Shape Among the phenotypic classes for thischaracter tall deltoid was dominant (31) followed by ovate(17) and narrow ovate (12) While local accessions inmost regions had tall deltoid bud shape 50 of the intro-duced accessions belong to two phenotypic classes roundwith central germpore (30) and 20 having pentagonalbud shape Equal proportions (19) of the samples fromBenshangul-Gumz were Ovate and Ovate with emarginatebasal wing
Canopy Structure Relative Plant Erectness Colour of theLeaves and Colour of the Exposed Rind Out of the eightphenotypic classes observed compact tip droopy was themost frequent (35) Open semidroopy and open droopyclasses had fairly equal distribution With regard to relativeplant erectness seventy-five percent of the germplasm werefound to fall in twophenotypic classes erect (41) and almost
erect (34) for the whole region and country Most of theaccessions (90) showed light green or green leaf colourbut 20 from introduced collections were greenish yellowAmong phenotypic classes for colour of the exposed rindbrownish yellow yellow brown and yellowish green were themost frequent with fairly equal distribution
Relative Shape of Dewlap Double crescent had larger fre-quency (22) while squarish descending and flaring typeof dewlap shape had fairly equal proportions Seventeenpercent of germplasm from Tigray region had squarishdeltoid dewlap shape
Relative Degree of Internode Alignment and Internode ShapeForty-four percent of the sample germplasm were slightlyzigzag while there was almost equal frequency for straightand zigzag internode alignment for the entire region andcountry Straight or nearly straight (slightly zigzag) internodealignment of cane stalks is a very important character formechanized farming and postharvest handling in sugarcane[51] Forty percent of the collections were observed havingconcave-convex type of internode shape while 29 wascylindrical and 14 conoidal Other classes had fairly equalproportion
Type of Auricle and Relative Shape of Ligule An equalproportion of the germplasm totally constituting 84 ofthe collection belong to four phenotypic classes namelythose with no auricle and others with transitional and shortand lanceolate auricles Sixteen percent of the collectionsalso showed equal frequency for the remaining classesBenshangul-Gumz and Tigray regions had each equal 13accessions with falcate type of auricle Sixty-seven percentof the collections belong to three ligule shape classes cres-cent with lozenge (27) broad-crescent (21) and linear-crescent (19) The germplasm had almost equal proportionfor the other phenotypic classes of this character Nineteenpercent of the accessions fromBenshangul-Gumz region haddeltoid shape of ligule
Stalk Corky Cracks Corky Patches Growth Cracks and BudGroove Eighty-nine percent of the germplasm across regionshad no stalk corky cracks with only eleven percent exhibitingthis character All the germplasm in Gambella Harari andSomali had no stalk corky cracks On the other hand eighty-one percent of the germplasm for the whole region showedthe presence of stalk corky patches All the local germplasmin Benshangul-Gumz Gambella and Tigray had stalk corkypatches whereas 75 of the germplasm from Harari Region67 from Somali Region and 24 from Oromia Regionhad no stalk corky patches In the entire region 75 ofthe germplasm had no stalk growth cracks with only 25showing this character Sixty-three percent of the germplasmwere found to have bud groove Howevermost of the samplesin Harari (75) Amhara (63) and Oromia (53) Regionshad no bud groove
The results of this study indicated the wide distributionof phenotypic classes for the characters considered whichindicates the existence of different races and combination
Advances in Agriculture 15
of races in local genotypes of different regions in EthiopiaThis diversity of germplasmbased onphenotypicmarkers canhelp sugarcane breeders in identifying the identity of geno-types as well as maintaining genetic diversity Differencesin morphological characters of different sugarcane varietieshave been reported [52] Piscitelli [53] showed importantexomorphological qualitative characters of sugarcane varietythat are not influenced by environmental factors and thus canbe used as a selection tool in any breeding program
The amount of phenotypic diversity estimates based onthe ShannonndashWeaver diversity index (H1015840) are shown inTable 9 The 16 characters differed in their distribution aswell as the amount of variation Individual traits showed adifferent pattern of variation among accessions Estimatesof H1015840 for individual characters varied from 049 for stalkcorky cracks to 100 for bud groove with an overall meanof 080 Most of the characters were highly polymorphicwhereas relative plant erectness and presence or absenceof stalk corky patches scored moderate values The highdiversity values for the characters showed a wide variabilityamong genotypes According to [54] this index is also usedin genetic resource studies as a convenient measure of bothallelic richness and allelic evenness when using geneticdata
The highest diversity computed was on presence orabsence of bud groove with a diversity index of 100indicating that this character is very variable in the localsugarcane germplasm On the other hand the descriptorcomputed with the lowest diversity index value (049) wasthe presence or absence of stalk corky cracks indicatingthat this character is selected in the development of thegenotypes In agreement with the present study [55] workingon S officinarum accessions from the world collection ofsugarcane germplasm obtained a low index of 025 and 035for presence or absence of stalk corky patches and cracksrespectively
Their report also showed high diversity index values forinternode alignment growth cracks internode shape budcushion bud extension bud groove and ligule shape Sim-ilarly [56] experimenting on field collections of sugarcaneaccessions in Philippine reported high ShannonndashWeaverdiversity for stalk growth cracks bud cushion stalk corkypatch stalk corky cracks bud extension and auricle outershape Unlike the present study they have reported high indexfor stalk corky cracksMedium index values were reported forstalk internode shape and low index values for ligule shapeand bud shape High variability of sugarcane varieties fordewlap shape was also reported in [57]
The H1015840 pooled across characters by region ranged from079 in Oromiya to 085 in Tigray with an overall aver-age of 081 (Table 8) Most of the regions showed highShannonndashWeaver diversity index The populations that hadthe highest H1015840 were that of Tigray and introduction The H1015840averaged over regions for different characters was found torange from 044 for presence or absence of stalk corky cracksto 094 for relative degree of internode alignment and budgroove with an overall average of 081 (Table 9)
Assessment of the ShannonndashWeaver index also showedsensitivity to both the number of descriptor classes and the
distribution within phenotypic classes in each region Amongregions the mean values for the ShannonndashWeaver index arenot significantly different but differences were found whenindividual characters were considered These observationssuggest that some regions have high diversity for particulartraits while in other regions selective pressures might havereduced the variation to certain genotypes
4 Conclusions
The large number of landraces observed during the cur-rent study demonstrates the existence of diverse geneticresources of sugarcane in Ethiopia The landraces wouldserve to broaden the genetic base of the available sugarcanegermplasm Generally distribution of the overall phenotypicdiversity among sugarcane landraces in Ethiopia is unevenOur results showed that there was a wide range of variationthat existed in the local sugarcane landraces studied both atregional and within region levels Selection and differenti-ation of types might have taken place along a geographicalpattern perhaps associated with climate and use The presentstudy contributes to updating sugarcane descriptors adoptedfrom USDA-ARS as well as Bioversity passport data forthe future collection and evaluation As Ethiopian SugarCorporation is in the course of launching breeding programthe observed substantial variation of landraces would enablesugarcane breeders to design and practice breeding andselection programs to improve the crop
Data Availability
The data used to support the findings of this study areavailable from the corresponding author upon request
Conflicts of Interest
The authors declare that they have no conflicts of interest
Acknowledgments
The authors are grateful for the financial grant of SugarCorporation of Ethiopia Many thanks are due to EthiopianBiodiversity Institute (EBI) for its technical advice duringcollection of the local sugarcane genotypes Sincere thanksare forwarded to Ethiopian smallholder farmers who pre-served sugarcane germplasm and shared their knowledgeand experience Regional Zonal and District (ldquoWeredardquo)Agriculture Bureaus across Ethiopia are acknowledged fortheir support during the study
Supplementary Materials
These supplementary materials contain different tables sup-porting the results of the study These include passport dataof sugarcane germplasm collected and related informationon the materials local sugarcane germplasm collected andused formorphological diversity study introduced sugarcane
16 Advances in Agriculture
Table9Sh
anno
n-WeaverD
iversityIndices(H1015840)a
ndmeanandsta
ndarderror(plusmnS
E)of
land
racesineach
region
basedon
16discretecharacters
Characters
Amha
raBe
nsha
ngul-G
umz
Introd
uced
Oromia
SNNP
Tigray
ELb
Total
Presence
orabsenceo
fbud
cushion(BUDCUSH
ION)
083
095
095
089
082
098
090
088
Relativ
edegreeo
fbud
extension(BUDEX
TEND)
077
079
093
074
072
087
080
078
Relativ
ebud
Shape(BU
DSH
APE
)078
096
089
081
085
078
084
078
Canop
ystructure(C
ANOPY
)089
098
095
081
078
096
089
084
Relativ
eshape
ofdewlap(D
EWLA
PSHAP)
087
094
092
090
082
096
090
086
Relativ
eplant
erectness(ER
ECT)
080
082
050
067
071
088
073
064
Relativ
edegreeo
finterno
dealignm
ent(IN
ALIGN)
096
076
098
097
098
100
094
098
Relativ
einterno
deshape(IN
SHAPE
)086
069
095
077
083
091
083
082
Color
oftheleaves(LE
AFC
OLO
R)087
071
090
064
074
071
076
076
Type
ofou
tera
uricle(A
URI
CLE
OUT)
080
092
087
079
083
093
086
082
Relativ
eshape
ofligule(LIGSH
APE
)086
094
082
085
090
083
087
083
Presence
orabsenceo
fstalkcorkyc
racks(ST
ALK
CORK
C)
044
054
035
055
051
025
044
049
Presence
orabsenceo
fstalkcorkyp
atches(STA
LKCORK
P)066
000
072
079
059
087
060
069
Presence
orabsenceo
fstalkgrow
thcracks
(STA
LKCRA
CK)
066
100
088
062
088
099
084
080
Color
ofthee
xposed
rind(R
INDCOLE
)085
096
089
084
086
076
086
085
Presence
orabsenceo
fbud
groo
ve(BUDGRO
OVE)
095
100
072
100
100
099
094
100
Means
080plusmn0
03
081plusmn0
06
083plusmn0
04
079plusmn0
03
080plusmn0
03
085plusmn0
05
081plusmn0
03
080
b EL=
averageo
fentire
locatio
n
Advances in Agriculture 17
varieties used for morphological diversity study and fre-quency distribution for 16 qualitative characters in sugarcaneby regions in Ethiopia and country of sources of introducedgenotypes (Supplementary Materials)
References
[1] ldquoSpreading the word about clean solutions from sugarcanrdquoAccessed 04072018 httpwwwsugarcaneorg
[2] Central Statistics Agency (CSA) ldquoAgricultural sample surveyArea and production of major crops private peasant holdingsrdquoStatistical Bulletin Addis Ababa Ethiopia 2017
[3] T Esayas Exploration and Collection Characterization GeneticDiversity Analysis and Association of Traits for Yield and YieldComponents of Sugarcane (Saccharumspp) in Ethiopia [MasterThesis] Haramaya University Dire Dawa Ethiopia 2014
[4] J R Harlan Crops and Man American Society of AgronomyCrop Science Society of America Madison WI USA 2ndedition 1992
[5] R W Allard Principles of Plant Breeding JohnWilley and SonsNew York NY USA 2nd edition 1999
[6] B T Roach and J Daniels ldquoA review of the origin and improve-ment of sugarcanerdquo in Proceedings of Coper sugar InternationalSugarcane Workshop pp 1ndash32 1987
[7] P P Patil S S Patil and D U Patel ldquoGenetic diversity in sug-arcane (Saccharum spp Complex)rdquo Journal of Pharmacognosyand Phytochemistry vol 6 no 6 pp 570ndash573 2017
[8] R B Singh B Singh andR K Singh ldquoStudy of genetic diversityof sugarcane (Saccharum) species and commercial varietiesthrough TRAP molecular markersrdquo Indian Journal of PlantPhysiology vol 22 no 3 pp 332ndash338 2017
[9] R W Allard ldquoPopulation structure and sampling methodsrdquo inGenetic Resources in PlantsmdashTheir Exploration and Conserva-tion O H Frankel and E Bennett Eds IBP Handbook no 11pp 97ndash107 FA Davis Company Philadelphia PA USA 1970
[10] D R Marshall and A H D Brown ldquoOptimum samplingstrategies in genetic conservationrdquo inCrop Genetic Resources forToday and Tomorrow O H Frankel and J G Hawkes Eds pp53ndash80 Cambridge University Press 1975
[11] J G Hawkes Crop Genetic Resources Field Collection ManualPudoc Wagnningen Netherland 1980
[12] J G Hawkes The Diversity of Crop Plants Harvard UniversityPress London England 1983
[13] B P K Yerima Manual for Good Laboratory Practice Foodand Agriculture Organization of United Nation Addis AbabaEthiopia 1993
[14] A Walkley and I A Black ldquoAn examination of the degtjareffmethod for determining soil organic matter and a proposedmodification of the chromic acid titrationmethodrdquo Soil Sciencevol 37 no 1 pp 29ndash38 1934
[15] S R Olsen and L E Sommers ldquoPhosphorusrdquo in AgronomyMonograph A L Page et al Ed Methods of soil analysis part2 pp 403ndash430 ASA and SSSAMadisonWI USA 2nd edition1982
[16] R H Bray and L T Kurtz ldquoDetermination of total organic andavailable forms of phosphorus in soilsrdquo Soil Science vol 59 no1 pp 39ndash46 1945
[17] S Sahlemedhin andB Taye ldquoProcedures for soil and plant anal-ysisrdquo Tech Rep 74 National Soil Research Center EthiopianAgricultural Research Organization 2000
[18] F C Stickler S Weaden and AW Pauli ldquoLeaf area determina-tion in grain sorghumrdquo Journal of Agronomy and Crop Sciencevol 53 pp 187-188 1961
[19] Munsell Color ChartThe Munsell Book of Color Matte Collec-tion Munsell Color Chart NewWindsor NY USA 1990
[20] H O Hartley ldquoThe maximum F-ratio as a short cut test forheterogeneity of variancesrdquo Biometrika vol 37 pp 308ndash3121950
[21] L Pecetti P Annicchiarico and A B Damania ldquoBiodiversity ina germplasm collection of durumwheatrdquo Euphytica vol 60 no3 pp 229ndash238 1992
[22] L Pecetti and A B Damania ldquoGeographic variation intetraploid wheat (Triticum turgidum ssp turgidum convardurum) landraces from two provinces in Ethiopiardquo GeneticResources and Crop Evolution vol 43 no 5 pp 395ndash407 1996
[23] S K Jain C O Qualset G M Bhatt and K K Wu ldquoGeo-graphical patterns of phenotypic diversity in a world collectionof durumwheatsrdquoCrop Science vol 15 no 5 pp 700ndash704 1975
[24] K K Nkongolo L Chinthu M Malusi and Z Vokhiwa ldquoPar-ticipatory variety selection and characterization of sorghum(Sorghum bicolour (L) Moench) elite accessions from Malaw-ian gene pool using farmer and breeder knowledgerdquo AfricanJournal of Agricultural Research vol 3 pp 273ndash283 2008
[25] M B Semira ldquoInnovative methods and metrics for agricul-ture and nutrition actions soil quality and human nutritionin Ethiopiardquo Leverhulme Center for Integrative Research onAgriculture and Health (LCIRAH) 2017
[26] I A Khan A Khatri M A Siddiqui G S Nizamani and SRaza ldquoPerformance of promising sugarcane clone for yield andquality traits in different ecological zones of Sindhirdquo PakistanJournal of Botany vol 36 no 1 pp 83ndash92 2004
[27] M F Perera M E Arias D Costilla et al ldquoGenetic diversityassessment and genotype identification in sugarcane based onDNAmarkers andmorphological traitsrdquo Euphytica vol 185 no3 pp 491ndash510 2012
[28] C G Neil M Kay and C C Jack ldquoDiversity among mainlandUSA sugarcane cultivars examined by SSR genotypingrdquo Journalof the American Society of Sugar Cane Technologists vol 29 pp36ndash52 2009
[29] S SchenckMWCrepeauK KWu P HMooreQ Yu andRMing ldquoGenetic diversity and relationships in native HawaiianSaccharum officinarum sugarcanerdquo Journal of Heredity vol 95no 4 pp 327ndash331 2004
[30] R Gebretsadik H Shimelis M D Laing P Tongoona andN Mandefro ldquoA diagnostic appraisal of the sorghum farmingsystem and breeding priorities in Striga infested agro-ecologiesof Ethiopiardquo Agricultural Systems vol 123 pp 54ndash61 2014
[31] L E V Cesar A T Bruzi J A R Nunes et al ldquoGeneticdivergence of sugarcane varieties collected in the region ofLavras Minas Gerais Brazilrdquo Genetics and Molecular Researchvol 14 no 4 pp 14043ndash14049 2015
[32] C Grenier P J Bramel J A Dahlberg et al ldquoSorghums of theSudan analysis of regional diversity and distributionrdquo GeneticResources and Crop Evolution vol 51 no 5 pp 489ndash500 2004
[33] N V Nair R Balakrishnan and T V Sreenivasan ldquoVariabilityfor quantitative traits in exotic hybrid germplasm of sugarcanerdquoGenetic Resources and Crop Evolution vol 45 no 5 pp 459ndash464 1998
[34] R Nagarajan S Alarmelu and R M Shanthi ldquoStudies onvariation in interspecific hybrids of Saccharumrdquo Sugar Tech vol2 no 3 pp 42ndash46 2000
18 Advances in Agriculture
[35] D N Kamat and J R P Singh ldquoVariability in Sugarcane underrain fed conditionrdquo Sugar Technology Short Communicationsvol 3 no 1-2 pp 65ndash67 2001
[36] B S Rishipal A S M Chaudhary and S P Kadian ldquoStudieson character interrelationship in clonal generation of sugarcane(Saccharum complex hybrids)rdquo Indian Sugar vol 47 no 11 pp907ndash911 1998
[37] S Thippeswamy S T Kajjidoni P M Salimath and J V GoudldquoCorrelation and path analysis for cane yield juice quality andtheir component traits in sugarcanerdquo Short Communication vol5 no 1-2 pp 65ndash72 2003
[38] P S Verma S Pal and N K Karma ldquoGenetic variability andcorrelation studies in sugarcanerdquo Indian Sugar vol 49 pp 125ndash128 1999
[39] H A Esechie J W Maranville and W M Ross ldquoRelationshipof stalk morphology and chemical composition to lodgingresistance in sorghumrdquo Crop Science vol 17 no 4 p 609 1977
[40] P K Das B C Jena and N Nayak ldquoCharacter association andpath analysis of sugar yield in sugarcanerdquo Indian Sugar vol 46no 10 pp 805ndash808 1997
[41] K J Sreekumar M Kuriakose M Thomas D Alexander andS Santhakumari ldquoVariability heritability and correlation onstudies on the yield and quality characters of sugarcanerdquo IndianSugar vol 44 no 4 pp 243ndash249 1994
[42] J R Harlan J M de Wet and E G Price ldquoComparativeevolution of cerealsrdquo Evolution vol 27 no 2 pp 311ndash325 1973
[43] A Elings ldquoDurum wheat landraces from syria II patterns ofvariationrdquo Euphytica vol 54 no 3 pp 231ndash243 1991
[44] K Tato ldquoRainfall in Ethiopiardquo Ethiopian Geographical Journalvol 2 pp 28ndash36 1964
[45] A B L Stemler J R Harlan and J M J deWet ldquoThe sorghumsof ethiopiardquo Economic Botany vol 31 no 4 pp 446ndash460 1977
[46] P Annicchiarico L Pecetti and A B Damania ldquoRelationshipsbetween phenotypic variation and climatic factors at collectingsites in durum wheat landracesrdquo Hereditas vol 122 no 2 pp163ndash167 1995
[47] A B Damania L Pecetti C O Qualset and B O HumeidldquoDiversity and geographic distribution of adaptive traits inTriticum turgidum L (durum group) wheat landraces fromTurkeyrdquoGenetic Resources and Crop Evolution vol 43 no 5 pp409ndash422 1996
[48] A Ayana and E Bekele ldquoGeographical patterns of morpho-logical variation in sorghum (Sorghum bicolor (L) Moench)germplasm from Ethiopia and Eritrea quantitative charactersrdquoEuphytica vol 115 no 2 pp 91ndash104 2000
[49] D S Falconer Introduction to Quantitative Genetics JohnWilley and Sons Inc New York NY USA 3rd edition 1993
[50] R S Thorpe ldquoBiochemical analysis of geographical variationand racial affinitiesrdquo Biology Review vol 51 pp 407ndash452 1976
[51] M Farooq ldquoMorphological and agricultural characteristicsof sugarcane in relation to mechanized agriculture at NARCIslamabadrdquo Portu Rico Journal vol 1 pp 51ndash61 1989
[52] M Akhtar N N Elahi and M Ashraf ldquoEvaluation of exoticsugarcane varieties for agronomic characters and productivityrdquoPakistan Journal of Biological Sciences vol 4 no 1 pp 37ndash402001
[53] F R Piscitelli ldquoPrincipals caracteres exomorfologicos de cul-tures de cana de azucarrdquo TUC Iindustraly Agricos vol 7 pp 49ndash57 1994 (Spanish)
[54] A H D Brown and B S Weir ldquoMeasuring genetic variability inplant populationsrdquo in Isozyme in Plant Genetics Breeding S DTanksley and T J Orton Eds part A Elsevier Science 1983
[55] R Balakrishnan N V Nair and T V Sreenivasan ldquoA methodfor establishing a core collection of Saccharum officinarum Lgermplasm based on quantitative-morphological datardquo GeneticResources and Crop Evolution vol 47 no 1 pp 1ndash9 2000
[56] C A Nestor and B Q Reynaldo Sugarcane Variety Improve-ment in Southeast Asia and the Pacific for Enhanced andSustainable Productivity ndash Germplasm and Disease IndexingComponents Institute of Plant Breeding (IPB) College ofAgriculture (CA) UP Los Banos (UPLB) 2003
[57] A Maqbool J Muhammad and A Sagheer ldquoAgronomic traitsand morphological characteristics of some exotic varieties ofsugarcanerdquoPakistan Journal of Agricultural Research vol 19 no4 2006
of races in local genotypes of different regions in EthiopiaThis diversity of germplasmbased onphenotypicmarkers canhelp sugarcane breeders in identifying the identity of geno-types as well as maintaining genetic diversity Differencesin morphological characters of different sugarcane varietieshave been reported [52] Piscitelli [53] showed importantexomorphological qualitative characters of sugarcane varietythat are not influenced by environmental factors and thus canbe used as a selection tool in any breeding program
The amount of phenotypic diversity estimates based onthe ShannonndashWeaver diversity index (H1015840) are shown inTable 9 The 16 characters differed in their distribution aswell as the amount of variation Individual traits showed adifferent pattern of variation among accessions Estimatesof H1015840 for individual characters varied from 049 for stalkcorky cracks to 100 for bud groove with an overall meanof 080 Most of the characters were highly polymorphicwhereas relative plant erectness and presence or absenceof stalk corky patches scored moderate values The highdiversity values for the characters showed a wide variabilityamong genotypes According to [54] this index is also usedin genetic resource studies as a convenient measure of bothallelic richness and allelic evenness when using geneticdata
The highest diversity computed was on presence orabsence of bud groove with a diversity index of 100indicating that this character is very variable in the localsugarcane germplasm On the other hand the descriptorcomputed with the lowest diversity index value (049) wasthe presence or absence of stalk corky cracks indicatingthat this character is selected in the development of thegenotypes In agreement with the present study [55] workingon S officinarum accessions from the world collection ofsugarcane germplasm obtained a low index of 025 and 035for presence or absence of stalk corky patches and cracksrespectively
Their report also showed high diversity index values forinternode alignment growth cracks internode shape budcushion bud extension bud groove and ligule shape Sim-ilarly [56] experimenting on field collections of sugarcaneaccessions in Philippine reported high ShannonndashWeaverdiversity for stalk growth cracks bud cushion stalk corkypatch stalk corky cracks bud extension and auricle outershape Unlike the present study they have reported high indexfor stalk corky cracksMedium index values were reported forstalk internode shape and low index values for ligule shapeand bud shape High variability of sugarcane varieties fordewlap shape was also reported in [57]
The H1015840 pooled across characters by region ranged from079 in Oromiya to 085 in Tigray with an overall aver-age of 081 (Table 8) Most of the regions showed highShannonndashWeaver diversity index The populations that hadthe highest H1015840 were that of Tigray and introduction The H1015840averaged over regions for different characters was found torange from 044 for presence or absence of stalk corky cracksto 094 for relative degree of internode alignment and budgroove with an overall average of 081 (Table 9)
Assessment of the ShannonndashWeaver index also showedsensitivity to both the number of descriptor classes and the
distribution within phenotypic classes in each region Amongregions the mean values for the ShannonndashWeaver index arenot significantly different but differences were found whenindividual characters were considered These observationssuggest that some regions have high diversity for particulartraits while in other regions selective pressures might havereduced the variation to certain genotypes
4 Conclusions
The large number of landraces observed during the cur-rent study demonstrates the existence of diverse geneticresources of sugarcane in Ethiopia The landraces wouldserve to broaden the genetic base of the available sugarcanegermplasm Generally distribution of the overall phenotypicdiversity among sugarcane landraces in Ethiopia is unevenOur results showed that there was a wide range of variationthat existed in the local sugarcane landraces studied both atregional and within region levels Selection and differenti-ation of types might have taken place along a geographicalpattern perhaps associated with climate and use The presentstudy contributes to updating sugarcane descriptors adoptedfrom USDA-ARS as well as Bioversity passport data forthe future collection and evaluation As Ethiopian SugarCorporation is in the course of launching breeding programthe observed substantial variation of landraces would enablesugarcane breeders to design and practice breeding andselection programs to improve the crop
Data Availability
The data used to support the findings of this study areavailable from the corresponding author upon request
Conflicts of Interest
The authors declare that they have no conflicts of interest
Acknowledgments
The authors are grateful for the financial grant of SugarCorporation of Ethiopia Many thanks are due to EthiopianBiodiversity Institute (EBI) for its technical advice duringcollection of the local sugarcane genotypes Sincere thanksare forwarded to Ethiopian smallholder farmers who pre-served sugarcane germplasm and shared their knowledgeand experience Regional Zonal and District (ldquoWeredardquo)Agriculture Bureaus across Ethiopia are acknowledged fortheir support during the study
Supplementary Materials
These supplementary materials contain different tables sup-porting the results of the study These include passport dataof sugarcane germplasm collected and related informationon the materials local sugarcane germplasm collected andused formorphological diversity study introduced sugarcane
16 Advances in Agriculture
Table9Sh
anno
n-WeaverD
iversityIndices(H1015840)a
ndmeanandsta
ndarderror(plusmnS
E)of
land
racesineach
region
basedon
16discretecharacters
Characters
Amha
raBe
nsha
ngul-G
umz
Introd
uced
Oromia
SNNP
Tigray
ELb
Total
Presence
orabsenceo
fbud
cushion(BUDCUSH
ION)
083
095
095
089
082
098
090
088
Relativ
edegreeo
fbud
extension(BUDEX
TEND)
077
079
093
074
072
087
080
078
Relativ
ebud
Shape(BU
DSH
APE
)078
096
089
081
085
078
084
078
Canop
ystructure(C
ANOPY
)089
098
095
081
078
096
089
084
Relativ
eshape
ofdewlap(D
EWLA
PSHAP)
087
094
092
090
082
096
090
086
Relativ
eplant
erectness(ER
ECT)
080
082
050
067
071
088
073
064
Relativ
edegreeo
finterno
dealignm
ent(IN
ALIGN)
096
076
098
097
098
100
094
098
Relativ
einterno
deshape(IN
SHAPE
)086
069
095
077
083
091
083
082
Color
oftheleaves(LE
AFC
OLO
R)087
071
090
064
074
071
076
076
Type
ofou
tera
uricle(A
URI
CLE
OUT)
080
092
087
079
083
093
086
082
Relativ
eshape
ofligule(LIGSH
APE
)086
094
082
085
090
083
087
083
Presence
orabsenceo
fstalkcorkyc
racks(ST
ALK
CORK
C)
044
054
035
055
051
025
044
049
Presence
orabsenceo
fstalkcorkyp
atches(STA
LKCORK
P)066
000
072
079
059
087
060
069
Presence
orabsenceo
fstalkgrow
thcracks
(STA
LKCRA
CK)
066
100
088
062
088
099
084
080
Color
ofthee
xposed
rind(R
INDCOLE
)085
096
089
084
086
076
086
085
Presence
orabsenceo
fbud
groo
ve(BUDGRO
OVE)
095
100
072
100
100
099
094
100
Means
080plusmn0
03
081plusmn0
06
083plusmn0
04
079plusmn0
03
080plusmn0
03
085plusmn0
05
081plusmn0
03
080
b EL=
averageo
fentire
locatio
n
Advances in Agriculture 17
varieties used for morphological diversity study and fre-quency distribution for 16 qualitative characters in sugarcaneby regions in Ethiopia and country of sources of introducedgenotypes (Supplementary Materials)
References
[1] ldquoSpreading the word about clean solutions from sugarcanrdquoAccessed 04072018 httpwwwsugarcaneorg
[2] Central Statistics Agency (CSA) ldquoAgricultural sample surveyArea and production of major crops private peasant holdingsrdquoStatistical Bulletin Addis Ababa Ethiopia 2017
[3] T Esayas Exploration and Collection Characterization GeneticDiversity Analysis and Association of Traits for Yield and YieldComponents of Sugarcane (Saccharumspp) in Ethiopia [MasterThesis] Haramaya University Dire Dawa Ethiopia 2014
[4] J R Harlan Crops and Man American Society of AgronomyCrop Science Society of America Madison WI USA 2ndedition 1992
[5] R W Allard Principles of Plant Breeding JohnWilley and SonsNew York NY USA 2nd edition 1999
[6] B T Roach and J Daniels ldquoA review of the origin and improve-ment of sugarcanerdquo in Proceedings of Coper sugar InternationalSugarcane Workshop pp 1ndash32 1987
[7] P P Patil S S Patil and D U Patel ldquoGenetic diversity in sug-arcane (Saccharum spp Complex)rdquo Journal of Pharmacognosyand Phytochemistry vol 6 no 6 pp 570ndash573 2017
[8] R B Singh B Singh andR K Singh ldquoStudy of genetic diversityof sugarcane (Saccharum) species and commercial varietiesthrough TRAP molecular markersrdquo Indian Journal of PlantPhysiology vol 22 no 3 pp 332ndash338 2017
[9] R W Allard ldquoPopulation structure and sampling methodsrdquo inGenetic Resources in PlantsmdashTheir Exploration and Conserva-tion O H Frankel and E Bennett Eds IBP Handbook no 11pp 97ndash107 FA Davis Company Philadelphia PA USA 1970
[10] D R Marshall and A H D Brown ldquoOptimum samplingstrategies in genetic conservationrdquo inCrop Genetic Resources forToday and Tomorrow O H Frankel and J G Hawkes Eds pp53ndash80 Cambridge University Press 1975
[11] J G Hawkes Crop Genetic Resources Field Collection ManualPudoc Wagnningen Netherland 1980
[12] J G Hawkes The Diversity of Crop Plants Harvard UniversityPress London England 1983
[13] B P K Yerima Manual for Good Laboratory Practice Foodand Agriculture Organization of United Nation Addis AbabaEthiopia 1993
[14] A Walkley and I A Black ldquoAn examination of the degtjareffmethod for determining soil organic matter and a proposedmodification of the chromic acid titrationmethodrdquo Soil Sciencevol 37 no 1 pp 29ndash38 1934
[15] S R Olsen and L E Sommers ldquoPhosphorusrdquo in AgronomyMonograph A L Page et al Ed Methods of soil analysis part2 pp 403ndash430 ASA and SSSAMadisonWI USA 2nd edition1982
[16] R H Bray and L T Kurtz ldquoDetermination of total organic andavailable forms of phosphorus in soilsrdquo Soil Science vol 59 no1 pp 39ndash46 1945
[17] S Sahlemedhin andB Taye ldquoProcedures for soil and plant anal-ysisrdquo Tech Rep 74 National Soil Research Center EthiopianAgricultural Research Organization 2000
[18] F C Stickler S Weaden and AW Pauli ldquoLeaf area determina-tion in grain sorghumrdquo Journal of Agronomy and Crop Sciencevol 53 pp 187-188 1961
[19] Munsell Color ChartThe Munsell Book of Color Matte Collec-tion Munsell Color Chart NewWindsor NY USA 1990
[20] H O Hartley ldquoThe maximum F-ratio as a short cut test forheterogeneity of variancesrdquo Biometrika vol 37 pp 308ndash3121950
[21] L Pecetti P Annicchiarico and A B Damania ldquoBiodiversity ina germplasm collection of durumwheatrdquo Euphytica vol 60 no3 pp 229ndash238 1992
[22] L Pecetti and A B Damania ldquoGeographic variation intetraploid wheat (Triticum turgidum ssp turgidum convardurum) landraces from two provinces in Ethiopiardquo GeneticResources and Crop Evolution vol 43 no 5 pp 395ndash407 1996
[23] S K Jain C O Qualset G M Bhatt and K K Wu ldquoGeo-graphical patterns of phenotypic diversity in a world collectionof durumwheatsrdquoCrop Science vol 15 no 5 pp 700ndash704 1975
[24] K K Nkongolo L Chinthu M Malusi and Z Vokhiwa ldquoPar-ticipatory variety selection and characterization of sorghum(Sorghum bicolour (L) Moench) elite accessions from Malaw-ian gene pool using farmer and breeder knowledgerdquo AfricanJournal of Agricultural Research vol 3 pp 273ndash283 2008
[25] M B Semira ldquoInnovative methods and metrics for agricul-ture and nutrition actions soil quality and human nutritionin Ethiopiardquo Leverhulme Center for Integrative Research onAgriculture and Health (LCIRAH) 2017
[26] I A Khan A Khatri M A Siddiqui G S Nizamani and SRaza ldquoPerformance of promising sugarcane clone for yield andquality traits in different ecological zones of Sindhirdquo PakistanJournal of Botany vol 36 no 1 pp 83ndash92 2004
[27] M F Perera M E Arias D Costilla et al ldquoGenetic diversityassessment and genotype identification in sugarcane based onDNAmarkers andmorphological traitsrdquo Euphytica vol 185 no3 pp 491ndash510 2012
[28] C G Neil M Kay and C C Jack ldquoDiversity among mainlandUSA sugarcane cultivars examined by SSR genotypingrdquo Journalof the American Society of Sugar Cane Technologists vol 29 pp36ndash52 2009
[29] S SchenckMWCrepeauK KWu P HMooreQ Yu andRMing ldquoGenetic diversity and relationships in native HawaiianSaccharum officinarum sugarcanerdquo Journal of Heredity vol 95no 4 pp 327ndash331 2004
[30] R Gebretsadik H Shimelis M D Laing P Tongoona andN Mandefro ldquoA diagnostic appraisal of the sorghum farmingsystem and breeding priorities in Striga infested agro-ecologiesof Ethiopiardquo Agricultural Systems vol 123 pp 54ndash61 2014
[31] L E V Cesar A T Bruzi J A R Nunes et al ldquoGeneticdivergence of sugarcane varieties collected in the region ofLavras Minas Gerais Brazilrdquo Genetics and Molecular Researchvol 14 no 4 pp 14043ndash14049 2015
[32] C Grenier P J Bramel J A Dahlberg et al ldquoSorghums of theSudan analysis of regional diversity and distributionrdquo GeneticResources and Crop Evolution vol 51 no 5 pp 489ndash500 2004
[33] N V Nair R Balakrishnan and T V Sreenivasan ldquoVariabilityfor quantitative traits in exotic hybrid germplasm of sugarcanerdquoGenetic Resources and Crop Evolution vol 45 no 5 pp 459ndash464 1998
[34] R Nagarajan S Alarmelu and R M Shanthi ldquoStudies onvariation in interspecific hybrids of Saccharumrdquo Sugar Tech vol2 no 3 pp 42ndash46 2000
18 Advances in Agriculture
[35] D N Kamat and J R P Singh ldquoVariability in Sugarcane underrain fed conditionrdquo Sugar Technology Short Communicationsvol 3 no 1-2 pp 65ndash67 2001
[36] B S Rishipal A S M Chaudhary and S P Kadian ldquoStudieson character interrelationship in clonal generation of sugarcane(Saccharum complex hybrids)rdquo Indian Sugar vol 47 no 11 pp907ndash911 1998
[37] S Thippeswamy S T Kajjidoni P M Salimath and J V GoudldquoCorrelation and path analysis for cane yield juice quality andtheir component traits in sugarcanerdquo Short Communication vol5 no 1-2 pp 65ndash72 2003
[38] P S Verma S Pal and N K Karma ldquoGenetic variability andcorrelation studies in sugarcanerdquo Indian Sugar vol 49 pp 125ndash128 1999
[39] H A Esechie J W Maranville and W M Ross ldquoRelationshipof stalk morphology and chemical composition to lodgingresistance in sorghumrdquo Crop Science vol 17 no 4 p 609 1977
[40] P K Das B C Jena and N Nayak ldquoCharacter association andpath analysis of sugar yield in sugarcanerdquo Indian Sugar vol 46no 10 pp 805ndash808 1997
[41] K J Sreekumar M Kuriakose M Thomas D Alexander andS Santhakumari ldquoVariability heritability and correlation onstudies on the yield and quality characters of sugarcanerdquo IndianSugar vol 44 no 4 pp 243ndash249 1994
[42] J R Harlan J M de Wet and E G Price ldquoComparativeevolution of cerealsrdquo Evolution vol 27 no 2 pp 311ndash325 1973
[43] A Elings ldquoDurum wheat landraces from syria II patterns ofvariationrdquo Euphytica vol 54 no 3 pp 231ndash243 1991
[44] K Tato ldquoRainfall in Ethiopiardquo Ethiopian Geographical Journalvol 2 pp 28ndash36 1964
[45] A B L Stemler J R Harlan and J M J deWet ldquoThe sorghumsof ethiopiardquo Economic Botany vol 31 no 4 pp 446ndash460 1977
[46] P Annicchiarico L Pecetti and A B Damania ldquoRelationshipsbetween phenotypic variation and climatic factors at collectingsites in durum wheat landracesrdquo Hereditas vol 122 no 2 pp163ndash167 1995
[47] A B Damania L Pecetti C O Qualset and B O HumeidldquoDiversity and geographic distribution of adaptive traits inTriticum turgidum L (durum group) wheat landraces fromTurkeyrdquoGenetic Resources and Crop Evolution vol 43 no 5 pp409ndash422 1996
[48] A Ayana and E Bekele ldquoGeographical patterns of morpho-logical variation in sorghum (Sorghum bicolor (L) Moench)germplasm from Ethiopia and Eritrea quantitative charactersrdquoEuphytica vol 115 no 2 pp 91ndash104 2000
[49] D S Falconer Introduction to Quantitative Genetics JohnWilley and Sons Inc New York NY USA 3rd edition 1993
[50] R S Thorpe ldquoBiochemical analysis of geographical variationand racial affinitiesrdquo Biology Review vol 51 pp 407ndash452 1976
[51] M Farooq ldquoMorphological and agricultural characteristicsof sugarcane in relation to mechanized agriculture at NARCIslamabadrdquo Portu Rico Journal vol 1 pp 51ndash61 1989
[52] M Akhtar N N Elahi and M Ashraf ldquoEvaluation of exoticsugarcane varieties for agronomic characters and productivityrdquoPakistan Journal of Biological Sciences vol 4 no 1 pp 37ndash402001
[53] F R Piscitelli ldquoPrincipals caracteres exomorfologicos de cul-tures de cana de azucarrdquo TUC Iindustraly Agricos vol 7 pp 49ndash57 1994 (Spanish)
[54] A H D Brown and B S Weir ldquoMeasuring genetic variability inplant populationsrdquo in Isozyme in Plant Genetics Breeding S DTanksley and T J Orton Eds part A Elsevier Science 1983
[55] R Balakrishnan N V Nair and T V Sreenivasan ldquoA methodfor establishing a core collection of Saccharum officinarum Lgermplasm based on quantitative-morphological datardquo GeneticResources and Crop Evolution vol 47 no 1 pp 1ndash9 2000
[56] C A Nestor and B Q Reynaldo Sugarcane Variety Improve-ment in Southeast Asia and the Pacific for Enhanced andSustainable Productivity ndash Germplasm and Disease IndexingComponents Institute of Plant Breeding (IPB) College ofAgriculture (CA) UP Los Banos (UPLB) 2003
[57] A Maqbool J Muhammad and A Sagheer ldquoAgronomic traitsand morphological characteristics of some exotic varieties ofsugarcanerdquoPakistan Journal of Agricultural Research vol 19 no4 2006
varieties used for morphological diversity study and fre-quency distribution for 16 qualitative characters in sugarcaneby regions in Ethiopia and country of sources of introducedgenotypes (Supplementary Materials)
References
[1] ldquoSpreading the word about clean solutions from sugarcanrdquoAccessed 04072018 httpwwwsugarcaneorg
[2] Central Statistics Agency (CSA) ldquoAgricultural sample surveyArea and production of major crops private peasant holdingsrdquoStatistical Bulletin Addis Ababa Ethiopia 2017
[3] T Esayas Exploration and Collection Characterization GeneticDiversity Analysis and Association of Traits for Yield and YieldComponents of Sugarcane (Saccharumspp) in Ethiopia [MasterThesis] Haramaya University Dire Dawa Ethiopia 2014
[4] J R Harlan Crops and Man American Society of AgronomyCrop Science Society of America Madison WI USA 2ndedition 1992
[5] R W Allard Principles of Plant Breeding JohnWilley and SonsNew York NY USA 2nd edition 1999
[6] B T Roach and J Daniels ldquoA review of the origin and improve-ment of sugarcanerdquo in Proceedings of Coper sugar InternationalSugarcane Workshop pp 1ndash32 1987
[7] P P Patil S S Patil and D U Patel ldquoGenetic diversity in sug-arcane (Saccharum spp Complex)rdquo Journal of Pharmacognosyand Phytochemistry vol 6 no 6 pp 570ndash573 2017
[8] R B Singh B Singh andR K Singh ldquoStudy of genetic diversityof sugarcane (Saccharum) species and commercial varietiesthrough TRAP molecular markersrdquo Indian Journal of PlantPhysiology vol 22 no 3 pp 332ndash338 2017
[9] R W Allard ldquoPopulation structure and sampling methodsrdquo inGenetic Resources in PlantsmdashTheir Exploration and Conserva-tion O H Frankel and E Bennett Eds IBP Handbook no 11pp 97ndash107 FA Davis Company Philadelphia PA USA 1970
[10] D R Marshall and A H D Brown ldquoOptimum samplingstrategies in genetic conservationrdquo inCrop Genetic Resources forToday and Tomorrow O H Frankel and J G Hawkes Eds pp53ndash80 Cambridge University Press 1975
[11] J G Hawkes Crop Genetic Resources Field Collection ManualPudoc Wagnningen Netherland 1980
[12] J G Hawkes The Diversity of Crop Plants Harvard UniversityPress London England 1983
[13] B P K Yerima Manual for Good Laboratory Practice Foodand Agriculture Organization of United Nation Addis AbabaEthiopia 1993
[14] A Walkley and I A Black ldquoAn examination of the degtjareffmethod for determining soil organic matter and a proposedmodification of the chromic acid titrationmethodrdquo Soil Sciencevol 37 no 1 pp 29ndash38 1934
[15] S R Olsen and L E Sommers ldquoPhosphorusrdquo in AgronomyMonograph A L Page et al Ed Methods of soil analysis part2 pp 403ndash430 ASA and SSSAMadisonWI USA 2nd edition1982
[16] R H Bray and L T Kurtz ldquoDetermination of total organic andavailable forms of phosphorus in soilsrdquo Soil Science vol 59 no1 pp 39ndash46 1945
[17] S Sahlemedhin andB Taye ldquoProcedures for soil and plant anal-ysisrdquo Tech Rep 74 National Soil Research Center EthiopianAgricultural Research Organization 2000
[18] F C Stickler S Weaden and AW Pauli ldquoLeaf area determina-tion in grain sorghumrdquo Journal of Agronomy and Crop Sciencevol 53 pp 187-188 1961
[19] Munsell Color ChartThe Munsell Book of Color Matte Collec-tion Munsell Color Chart NewWindsor NY USA 1990
[20] H O Hartley ldquoThe maximum F-ratio as a short cut test forheterogeneity of variancesrdquo Biometrika vol 37 pp 308ndash3121950
[21] L Pecetti P Annicchiarico and A B Damania ldquoBiodiversity ina germplasm collection of durumwheatrdquo Euphytica vol 60 no3 pp 229ndash238 1992
[22] L Pecetti and A B Damania ldquoGeographic variation intetraploid wheat (Triticum turgidum ssp turgidum convardurum) landraces from two provinces in Ethiopiardquo GeneticResources and Crop Evolution vol 43 no 5 pp 395ndash407 1996
[23] S K Jain C O Qualset G M Bhatt and K K Wu ldquoGeo-graphical patterns of phenotypic diversity in a world collectionof durumwheatsrdquoCrop Science vol 15 no 5 pp 700ndash704 1975
[24] K K Nkongolo L Chinthu M Malusi and Z Vokhiwa ldquoPar-ticipatory variety selection and characterization of sorghum(Sorghum bicolour (L) Moench) elite accessions from Malaw-ian gene pool using farmer and breeder knowledgerdquo AfricanJournal of Agricultural Research vol 3 pp 273ndash283 2008
[25] M B Semira ldquoInnovative methods and metrics for agricul-ture and nutrition actions soil quality and human nutritionin Ethiopiardquo Leverhulme Center for Integrative Research onAgriculture and Health (LCIRAH) 2017
[26] I A Khan A Khatri M A Siddiqui G S Nizamani and SRaza ldquoPerformance of promising sugarcane clone for yield andquality traits in different ecological zones of Sindhirdquo PakistanJournal of Botany vol 36 no 1 pp 83ndash92 2004
[27] M F Perera M E Arias D Costilla et al ldquoGenetic diversityassessment and genotype identification in sugarcane based onDNAmarkers andmorphological traitsrdquo Euphytica vol 185 no3 pp 491ndash510 2012
[28] C G Neil M Kay and C C Jack ldquoDiversity among mainlandUSA sugarcane cultivars examined by SSR genotypingrdquo Journalof the American Society of Sugar Cane Technologists vol 29 pp36ndash52 2009
[29] S SchenckMWCrepeauK KWu P HMooreQ Yu andRMing ldquoGenetic diversity and relationships in native HawaiianSaccharum officinarum sugarcanerdquo Journal of Heredity vol 95no 4 pp 327ndash331 2004
[30] R Gebretsadik H Shimelis M D Laing P Tongoona andN Mandefro ldquoA diagnostic appraisal of the sorghum farmingsystem and breeding priorities in Striga infested agro-ecologiesof Ethiopiardquo Agricultural Systems vol 123 pp 54ndash61 2014
[31] L E V Cesar A T Bruzi J A R Nunes et al ldquoGeneticdivergence of sugarcane varieties collected in the region ofLavras Minas Gerais Brazilrdquo Genetics and Molecular Researchvol 14 no 4 pp 14043ndash14049 2015
[32] C Grenier P J Bramel J A Dahlberg et al ldquoSorghums of theSudan analysis of regional diversity and distributionrdquo GeneticResources and Crop Evolution vol 51 no 5 pp 489ndash500 2004
[33] N V Nair R Balakrishnan and T V Sreenivasan ldquoVariabilityfor quantitative traits in exotic hybrid germplasm of sugarcanerdquoGenetic Resources and Crop Evolution vol 45 no 5 pp 459ndash464 1998
[34] R Nagarajan S Alarmelu and R M Shanthi ldquoStudies onvariation in interspecific hybrids of Saccharumrdquo Sugar Tech vol2 no 3 pp 42ndash46 2000
18 Advances in Agriculture
[35] D N Kamat and J R P Singh ldquoVariability in Sugarcane underrain fed conditionrdquo Sugar Technology Short Communicationsvol 3 no 1-2 pp 65ndash67 2001
[36] B S Rishipal A S M Chaudhary and S P Kadian ldquoStudieson character interrelationship in clonal generation of sugarcane(Saccharum complex hybrids)rdquo Indian Sugar vol 47 no 11 pp907ndash911 1998
[37] S Thippeswamy S T Kajjidoni P M Salimath and J V GoudldquoCorrelation and path analysis for cane yield juice quality andtheir component traits in sugarcanerdquo Short Communication vol5 no 1-2 pp 65ndash72 2003
[38] P S Verma S Pal and N K Karma ldquoGenetic variability andcorrelation studies in sugarcanerdquo Indian Sugar vol 49 pp 125ndash128 1999
[39] H A Esechie J W Maranville and W M Ross ldquoRelationshipof stalk morphology and chemical composition to lodgingresistance in sorghumrdquo Crop Science vol 17 no 4 p 609 1977
[40] P K Das B C Jena and N Nayak ldquoCharacter association andpath analysis of sugar yield in sugarcanerdquo Indian Sugar vol 46no 10 pp 805ndash808 1997
[41] K J Sreekumar M Kuriakose M Thomas D Alexander andS Santhakumari ldquoVariability heritability and correlation onstudies on the yield and quality characters of sugarcanerdquo IndianSugar vol 44 no 4 pp 243ndash249 1994
[42] J R Harlan J M de Wet and E G Price ldquoComparativeevolution of cerealsrdquo Evolution vol 27 no 2 pp 311ndash325 1973
[43] A Elings ldquoDurum wheat landraces from syria II patterns ofvariationrdquo Euphytica vol 54 no 3 pp 231ndash243 1991
[44] K Tato ldquoRainfall in Ethiopiardquo Ethiopian Geographical Journalvol 2 pp 28ndash36 1964
[45] A B L Stemler J R Harlan and J M J deWet ldquoThe sorghumsof ethiopiardquo Economic Botany vol 31 no 4 pp 446ndash460 1977
[46] P Annicchiarico L Pecetti and A B Damania ldquoRelationshipsbetween phenotypic variation and climatic factors at collectingsites in durum wheat landracesrdquo Hereditas vol 122 no 2 pp163ndash167 1995
[47] A B Damania L Pecetti C O Qualset and B O HumeidldquoDiversity and geographic distribution of adaptive traits inTriticum turgidum L (durum group) wheat landraces fromTurkeyrdquoGenetic Resources and Crop Evolution vol 43 no 5 pp409ndash422 1996
[48] A Ayana and E Bekele ldquoGeographical patterns of morpho-logical variation in sorghum (Sorghum bicolor (L) Moench)germplasm from Ethiopia and Eritrea quantitative charactersrdquoEuphytica vol 115 no 2 pp 91ndash104 2000
[49] D S Falconer Introduction to Quantitative Genetics JohnWilley and Sons Inc New York NY USA 3rd edition 1993
[50] R S Thorpe ldquoBiochemical analysis of geographical variationand racial affinitiesrdquo Biology Review vol 51 pp 407ndash452 1976
[51] M Farooq ldquoMorphological and agricultural characteristicsof sugarcane in relation to mechanized agriculture at NARCIslamabadrdquo Portu Rico Journal vol 1 pp 51ndash61 1989
[52] M Akhtar N N Elahi and M Ashraf ldquoEvaluation of exoticsugarcane varieties for agronomic characters and productivityrdquoPakistan Journal of Biological Sciences vol 4 no 1 pp 37ndash402001
[53] F R Piscitelli ldquoPrincipals caracteres exomorfologicos de cul-tures de cana de azucarrdquo TUC Iindustraly Agricos vol 7 pp 49ndash57 1994 (Spanish)
[54] A H D Brown and B S Weir ldquoMeasuring genetic variability inplant populationsrdquo in Isozyme in Plant Genetics Breeding S DTanksley and T J Orton Eds part A Elsevier Science 1983
[55] R Balakrishnan N V Nair and T V Sreenivasan ldquoA methodfor establishing a core collection of Saccharum officinarum Lgermplasm based on quantitative-morphological datardquo GeneticResources and Crop Evolution vol 47 no 1 pp 1ndash9 2000
[56] C A Nestor and B Q Reynaldo Sugarcane Variety Improve-ment in Southeast Asia and the Pacific for Enhanced andSustainable Productivity ndash Germplasm and Disease IndexingComponents Institute of Plant Breeding (IPB) College ofAgriculture (CA) UP Los Banos (UPLB) 2003
[57] A Maqbool J Muhammad and A Sagheer ldquoAgronomic traitsand morphological characteristics of some exotic varieties ofsugarcanerdquoPakistan Journal of Agricultural Research vol 19 no4 2006
varieties used for morphological diversity study and fre-quency distribution for 16 qualitative characters in sugarcaneby regions in Ethiopia and country of sources of introducedgenotypes (Supplementary Materials)
References
[1] ldquoSpreading the word about clean solutions from sugarcanrdquoAccessed 04072018 httpwwwsugarcaneorg
[2] Central Statistics Agency (CSA) ldquoAgricultural sample surveyArea and production of major crops private peasant holdingsrdquoStatistical Bulletin Addis Ababa Ethiopia 2017
[3] T Esayas Exploration and Collection Characterization GeneticDiversity Analysis and Association of Traits for Yield and YieldComponents of Sugarcane (Saccharumspp) in Ethiopia [MasterThesis] Haramaya University Dire Dawa Ethiopia 2014
[4] J R Harlan Crops and Man American Society of AgronomyCrop Science Society of America Madison WI USA 2ndedition 1992
[5] R W Allard Principles of Plant Breeding JohnWilley and SonsNew York NY USA 2nd edition 1999
[6] B T Roach and J Daniels ldquoA review of the origin and improve-ment of sugarcanerdquo in Proceedings of Coper sugar InternationalSugarcane Workshop pp 1ndash32 1987
[7] P P Patil S S Patil and D U Patel ldquoGenetic diversity in sug-arcane (Saccharum spp Complex)rdquo Journal of Pharmacognosyand Phytochemistry vol 6 no 6 pp 570ndash573 2017
[8] R B Singh B Singh andR K Singh ldquoStudy of genetic diversityof sugarcane (Saccharum) species and commercial varietiesthrough TRAP molecular markersrdquo Indian Journal of PlantPhysiology vol 22 no 3 pp 332ndash338 2017
[9] R W Allard ldquoPopulation structure and sampling methodsrdquo inGenetic Resources in PlantsmdashTheir Exploration and Conserva-tion O H Frankel and E Bennett Eds IBP Handbook no 11pp 97ndash107 FA Davis Company Philadelphia PA USA 1970
[10] D R Marshall and A H D Brown ldquoOptimum samplingstrategies in genetic conservationrdquo inCrop Genetic Resources forToday and Tomorrow O H Frankel and J G Hawkes Eds pp53ndash80 Cambridge University Press 1975
[11] J G Hawkes Crop Genetic Resources Field Collection ManualPudoc Wagnningen Netherland 1980
[12] J G Hawkes The Diversity of Crop Plants Harvard UniversityPress London England 1983
[13] B P K Yerima Manual for Good Laboratory Practice Foodand Agriculture Organization of United Nation Addis AbabaEthiopia 1993
[14] A Walkley and I A Black ldquoAn examination of the degtjareffmethod for determining soil organic matter and a proposedmodification of the chromic acid titrationmethodrdquo Soil Sciencevol 37 no 1 pp 29ndash38 1934
[15] S R Olsen and L E Sommers ldquoPhosphorusrdquo in AgronomyMonograph A L Page et al Ed Methods of soil analysis part2 pp 403ndash430 ASA and SSSAMadisonWI USA 2nd edition1982
[16] R H Bray and L T Kurtz ldquoDetermination of total organic andavailable forms of phosphorus in soilsrdquo Soil Science vol 59 no1 pp 39ndash46 1945
[17] S Sahlemedhin andB Taye ldquoProcedures for soil and plant anal-ysisrdquo Tech Rep 74 National Soil Research Center EthiopianAgricultural Research Organization 2000
[18] F C Stickler S Weaden and AW Pauli ldquoLeaf area determina-tion in grain sorghumrdquo Journal of Agronomy and Crop Sciencevol 53 pp 187-188 1961
[19] Munsell Color ChartThe Munsell Book of Color Matte Collec-tion Munsell Color Chart NewWindsor NY USA 1990
[20] H O Hartley ldquoThe maximum F-ratio as a short cut test forheterogeneity of variancesrdquo Biometrika vol 37 pp 308ndash3121950
[21] L Pecetti P Annicchiarico and A B Damania ldquoBiodiversity ina germplasm collection of durumwheatrdquo Euphytica vol 60 no3 pp 229ndash238 1992
[22] L Pecetti and A B Damania ldquoGeographic variation intetraploid wheat (Triticum turgidum ssp turgidum convardurum) landraces from two provinces in Ethiopiardquo GeneticResources and Crop Evolution vol 43 no 5 pp 395ndash407 1996
[23] S K Jain C O Qualset G M Bhatt and K K Wu ldquoGeo-graphical patterns of phenotypic diversity in a world collectionof durumwheatsrdquoCrop Science vol 15 no 5 pp 700ndash704 1975
[24] K K Nkongolo L Chinthu M Malusi and Z Vokhiwa ldquoPar-ticipatory variety selection and characterization of sorghum(Sorghum bicolour (L) Moench) elite accessions from Malaw-ian gene pool using farmer and breeder knowledgerdquo AfricanJournal of Agricultural Research vol 3 pp 273ndash283 2008
[25] M B Semira ldquoInnovative methods and metrics for agricul-ture and nutrition actions soil quality and human nutritionin Ethiopiardquo Leverhulme Center for Integrative Research onAgriculture and Health (LCIRAH) 2017
[26] I A Khan A Khatri M A Siddiqui G S Nizamani and SRaza ldquoPerformance of promising sugarcane clone for yield andquality traits in different ecological zones of Sindhirdquo PakistanJournal of Botany vol 36 no 1 pp 83ndash92 2004
[27] M F Perera M E Arias D Costilla et al ldquoGenetic diversityassessment and genotype identification in sugarcane based onDNAmarkers andmorphological traitsrdquo Euphytica vol 185 no3 pp 491ndash510 2012
[28] C G Neil M Kay and C C Jack ldquoDiversity among mainlandUSA sugarcane cultivars examined by SSR genotypingrdquo Journalof the American Society of Sugar Cane Technologists vol 29 pp36ndash52 2009
[29] S SchenckMWCrepeauK KWu P HMooreQ Yu andRMing ldquoGenetic diversity and relationships in native HawaiianSaccharum officinarum sugarcanerdquo Journal of Heredity vol 95no 4 pp 327ndash331 2004
[30] R Gebretsadik H Shimelis M D Laing P Tongoona andN Mandefro ldquoA diagnostic appraisal of the sorghum farmingsystem and breeding priorities in Striga infested agro-ecologiesof Ethiopiardquo Agricultural Systems vol 123 pp 54ndash61 2014
[31] L E V Cesar A T Bruzi J A R Nunes et al ldquoGeneticdivergence of sugarcane varieties collected in the region ofLavras Minas Gerais Brazilrdquo Genetics and Molecular Researchvol 14 no 4 pp 14043ndash14049 2015
[32] C Grenier P J Bramel J A Dahlberg et al ldquoSorghums of theSudan analysis of regional diversity and distributionrdquo GeneticResources and Crop Evolution vol 51 no 5 pp 489ndash500 2004
[33] N V Nair R Balakrishnan and T V Sreenivasan ldquoVariabilityfor quantitative traits in exotic hybrid germplasm of sugarcanerdquoGenetic Resources and Crop Evolution vol 45 no 5 pp 459ndash464 1998
[34] R Nagarajan S Alarmelu and R M Shanthi ldquoStudies onvariation in interspecific hybrids of Saccharumrdquo Sugar Tech vol2 no 3 pp 42ndash46 2000
18 Advances in Agriculture
[35] D N Kamat and J R P Singh ldquoVariability in Sugarcane underrain fed conditionrdquo Sugar Technology Short Communicationsvol 3 no 1-2 pp 65ndash67 2001
[36] B S Rishipal A S M Chaudhary and S P Kadian ldquoStudieson character interrelationship in clonal generation of sugarcane(Saccharum complex hybrids)rdquo Indian Sugar vol 47 no 11 pp907ndash911 1998
[37] S Thippeswamy S T Kajjidoni P M Salimath and J V GoudldquoCorrelation and path analysis for cane yield juice quality andtheir component traits in sugarcanerdquo Short Communication vol5 no 1-2 pp 65ndash72 2003
[38] P S Verma S Pal and N K Karma ldquoGenetic variability andcorrelation studies in sugarcanerdquo Indian Sugar vol 49 pp 125ndash128 1999
[39] H A Esechie J W Maranville and W M Ross ldquoRelationshipof stalk morphology and chemical composition to lodgingresistance in sorghumrdquo Crop Science vol 17 no 4 p 609 1977
[40] P K Das B C Jena and N Nayak ldquoCharacter association andpath analysis of sugar yield in sugarcanerdquo Indian Sugar vol 46no 10 pp 805ndash808 1997
[41] K J Sreekumar M Kuriakose M Thomas D Alexander andS Santhakumari ldquoVariability heritability and correlation onstudies on the yield and quality characters of sugarcanerdquo IndianSugar vol 44 no 4 pp 243ndash249 1994
[42] J R Harlan J M de Wet and E G Price ldquoComparativeevolution of cerealsrdquo Evolution vol 27 no 2 pp 311ndash325 1973
[43] A Elings ldquoDurum wheat landraces from syria II patterns ofvariationrdquo Euphytica vol 54 no 3 pp 231ndash243 1991
[44] K Tato ldquoRainfall in Ethiopiardquo Ethiopian Geographical Journalvol 2 pp 28ndash36 1964
[45] A B L Stemler J R Harlan and J M J deWet ldquoThe sorghumsof ethiopiardquo Economic Botany vol 31 no 4 pp 446ndash460 1977
[46] P Annicchiarico L Pecetti and A B Damania ldquoRelationshipsbetween phenotypic variation and climatic factors at collectingsites in durum wheat landracesrdquo Hereditas vol 122 no 2 pp163ndash167 1995
[47] A B Damania L Pecetti C O Qualset and B O HumeidldquoDiversity and geographic distribution of adaptive traits inTriticum turgidum L (durum group) wheat landraces fromTurkeyrdquoGenetic Resources and Crop Evolution vol 43 no 5 pp409ndash422 1996
[48] A Ayana and E Bekele ldquoGeographical patterns of morpho-logical variation in sorghum (Sorghum bicolor (L) Moench)germplasm from Ethiopia and Eritrea quantitative charactersrdquoEuphytica vol 115 no 2 pp 91ndash104 2000
[49] D S Falconer Introduction to Quantitative Genetics JohnWilley and Sons Inc New York NY USA 3rd edition 1993
[50] R S Thorpe ldquoBiochemical analysis of geographical variationand racial affinitiesrdquo Biology Review vol 51 pp 407ndash452 1976
[51] M Farooq ldquoMorphological and agricultural characteristicsof sugarcane in relation to mechanized agriculture at NARCIslamabadrdquo Portu Rico Journal vol 1 pp 51ndash61 1989
[52] M Akhtar N N Elahi and M Ashraf ldquoEvaluation of exoticsugarcane varieties for agronomic characters and productivityrdquoPakistan Journal of Biological Sciences vol 4 no 1 pp 37ndash402001
[53] F R Piscitelli ldquoPrincipals caracteres exomorfologicos de cul-tures de cana de azucarrdquo TUC Iindustraly Agricos vol 7 pp 49ndash57 1994 (Spanish)
[54] A H D Brown and B S Weir ldquoMeasuring genetic variability inplant populationsrdquo in Isozyme in Plant Genetics Breeding S DTanksley and T J Orton Eds part A Elsevier Science 1983
[55] R Balakrishnan N V Nair and T V Sreenivasan ldquoA methodfor establishing a core collection of Saccharum officinarum Lgermplasm based on quantitative-morphological datardquo GeneticResources and Crop Evolution vol 47 no 1 pp 1ndash9 2000
[56] C A Nestor and B Q Reynaldo Sugarcane Variety Improve-ment in Southeast Asia and the Pacific for Enhanced andSustainable Productivity ndash Germplasm and Disease IndexingComponents Institute of Plant Breeding (IPB) College ofAgriculture (CA) UP Los Banos (UPLB) 2003
[57] A Maqbool J Muhammad and A Sagheer ldquoAgronomic traitsand morphological characteristics of some exotic varieties ofsugarcanerdquoPakistan Journal of Agricultural Research vol 19 no4 2006
[35] D N Kamat and J R P Singh ldquoVariability in Sugarcane underrain fed conditionrdquo Sugar Technology Short Communicationsvol 3 no 1-2 pp 65ndash67 2001
[36] B S Rishipal A S M Chaudhary and S P Kadian ldquoStudieson character interrelationship in clonal generation of sugarcane(Saccharum complex hybrids)rdquo Indian Sugar vol 47 no 11 pp907ndash911 1998
[37] S Thippeswamy S T Kajjidoni P M Salimath and J V GoudldquoCorrelation and path analysis for cane yield juice quality andtheir component traits in sugarcanerdquo Short Communication vol5 no 1-2 pp 65ndash72 2003
[38] P S Verma S Pal and N K Karma ldquoGenetic variability andcorrelation studies in sugarcanerdquo Indian Sugar vol 49 pp 125ndash128 1999
[39] H A Esechie J W Maranville and W M Ross ldquoRelationshipof stalk morphology and chemical composition to lodgingresistance in sorghumrdquo Crop Science vol 17 no 4 p 609 1977
[40] P K Das B C Jena and N Nayak ldquoCharacter association andpath analysis of sugar yield in sugarcanerdquo Indian Sugar vol 46no 10 pp 805ndash808 1997
[41] K J Sreekumar M Kuriakose M Thomas D Alexander andS Santhakumari ldquoVariability heritability and correlation onstudies on the yield and quality characters of sugarcanerdquo IndianSugar vol 44 no 4 pp 243ndash249 1994
[42] J R Harlan J M de Wet and E G Price ldquoComparativeevolution of cerealsrdquo Evolution vol 27 no 2 pp 311ndash325 1973
[43] A Elings ldquoDurum wheat landraces from syria II patterns ofvariationrdquo Euphytica vol 54 no 3 pp 231ndash243 1991
[44] K Tato ldquoRainfall in Ethiopiardquo Ethiopian Geographical Journalvol 2 pp 28ndash36 1964
[45] A B L Stemler J R Harlan and J M J deWet ldquoThe sorghumsof ethiopiardquo Economic Botany vol 31 no 4 pp 446ndash460 1977
[46] P Annicchiarico L Pecetti and A B Damania ldquoRelationshipsbetween phenotypic variation and climatic factors at collectingsites in durum wheat landracesrdquo Hereditas vol 122 no 2 pp163ndash167 1995
[47] A B Damania L Pecetti C O Qualset and B O HumeidldquoDiversity and geographic distribution of adaptive traits inTriticum turgidum L (durum group) wheat landraces fromTurkeyrdquoGenetic Resources and Crop Evolution vol 43 no 5 pp409ndash422 1996
[48] A Ayana and E Bekele ldquoGeographical patterns of morpho-logical variation in sorghum (Sorghum bicolor (L) Moench)germplasm from Ethiopia and Eritrea quantitative charactersrdquoEuphytica vol 115 no 2 pp 91ndash104 2000
[49] D S Falconer Introduction to Quantitative Genetics JohnWilley and Sons Inc New York NY USA 3rd edition 1993
[50] R S Thorpe ldquoBiochemical analysis of geographical variationand racial affinitiesrdquo Biology Review vol 51 pp 407ndash452 1976
[51] M Farooq ldquoMorphological and agricultural characteristicsof sugarcane in relation to mechanized agriculture at NARCIslamabadrdquo Portu Rico Journal vol 1 pp 51ndash61 1989
[52] M Akhtar N N Elahi and M Ashraf ldquoEvaluation of exoticsugarcane varieties for agronomic characters and productivityrdquoPakistan Journal of Biological Sciences vol 4 no 1 pp 37ndash402001
[53] F R Piscitelli ldquoPrincipals caracteres exomorfologicos de cul-tures de cana de azucarrdquo TUC Iindustraly Agricos vol 7 pp 49ndash57 1994 (Spanish)
[54] A H D Brown and B S Weir ldquoMeasuring genetic variability inplant populationsrdquo in Isozyme in Plant Genetics Breeding S DTanksley and T J Orton Eds part A Elsevier Science 1983
[55] R Balakrishnan N V Nair and T V Sreenivasan ldquoA methodfor establishing a core collection of Saccharum officinarum Lgermplasm based on quantitative-morphological datardquo GeneticResources and Crop Evolution vol 47 no 1 pp 1ndash9 2000
[56] C A Nestor and B Q Reynaldo Sugarcane Variety Improve-ment in Southeast Asia and the Pacific for Enhanced andSustainable Productivity ndash Germplasm and Disease IndexingComponents Institute of Plant Breeding (IPB) College ofAgriculture (CA) UP Los Banos (UPLB) 2003
[57] A Maqbool J Muhammad and A Sagheer ldquoAgronomic traitsand morphological characteristics of some exotic varieties ofsugarcanerdquoPakistan Journal of Agricultural Research vol 19 no4 2006