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Inducing drought tolerance in plants Recent advances
M Ashraf
Department of Botany University of Agriculture Faisalabad Pakistan
a b s t r a c ta r t i c l e i n f o
Article history
Received 21 September 2009
Received in revised form 6 November 2009
Accepted 9 November 2009
Available online 13 November 2009
Keywords
Drought tolerance
Conventional breeding
Marker assisted selection
Quantitative trait loci
Transgenic plants
Osmolytes
Hormones
Recombinant inbred lines
Undoubtedly drought is one of the prime abiotic stresses in the world Crop yield losses due to drought
stress are considerable Although a variety of approaches have been used to alleviate the problem of drought
plant breeding either conventional breeding or genetic engineering seems to be an ef 1047297cient and economic
means of tailoring crops to enable them to grow successfully in drought-prone environments During the lastcentury although plant breeders have made ample progress through conventional breeding in developing
drought tolerant linescultivars of some selected crops the approach is in fact highly time-consuming and
labor- and cost-intensive Alternatively marker-assisted breeding (MAB) is a more ef 1047297cient approach which
identi1047297es the usefulness of thousands of genomic regions of a crop under stress conditions which was in
reality previously not possible Quantitative trait loci (QTL) for drought tolerance have been identi 1047297ed for a
variety of traits in different crops With the development of comprehensive molecular linkage maps marker-
assisted selection procedures have led to pyramiding desirable traits to achieve improvements in crop
drought tolerance However the accuracy and preciseness in QTL identi1047297cation are problematic
Furthermore signi1047297cant genetictimesenvironment interaction large number of genes encoding yield and use
of wrong mapping populations have all harmed programs involved in mapping of QTL for high growth and
yield under water limited conditions Under such circumstances a transgenic approach to the problem
seems more convincing and practicable and it is being pursued vigorously to improve qualitative and
quantitative traits including tolerance to biotic and abiotic stresses in different crops Rapid advance in
knowledge on genomics and proteomics will certainly be bene1047297cial to 1047297ne-tune the molecular breeding and
transformation approaches so as to achieve a signi1047297cant progress in crop improvement in future Knowledge
of gene regulation and signal transduction to generate drought tolerant crop cultivarslines has beendiscussed in the present review In addition the advantages and disadvantages as well as future prospects of
each breeding approach have also been discussed
copy 2009 Elsevier Inc All rights reserved
Contents
1 Introduction 169
2 Conventional breeding for drought tolerance 170
3 Marker-assisted breeding (MAB) for drought tolerance 171
31 Identi1047297cation of QTL associated with drought tolerance 171
32 Manipulation of QTL for developing drought tolerant crop cultivarslines 173
4 Engineering crops for enhanced drought tolerancemdashTransgenic approach 176
5 Conclusions and future challenges 180References 181
1 Introduction
Plant breeding conventional breeding or genetic engineering is an
art through which crop varietiesof high quality and yield are developed
Breeding forany desired trait undoubtedly requires a signi1047297cant amount
of genetic variation at intra-speci1047297c inter-speci1047297c or inter-generic levels
Variability however can be achieved by new gene combinations
intercrossing those genotypes that hold desirable characteristics and
introducing new germplasm from other existing breeding programs
(Ashraf 1994 Flowers 2004 Ashraf and Akram 2009) The variability
achieved through these means canbe narrowed down by selectinga few
Biotechnology Advances 28 (2010) 169ndash183
E-mail address ashrafbotyahoocom
0734-9750$ ndash see front matter copy 2009 Elsevier Inc All rights reserved
doi101016jbiotechadv200911005
Contents lists available at ScienceDirect
Biotechnology Advances
j o u r n a l h o m e p a g e w w w e l s ev i e r c o m l o c a t e b i o t e c h a d v
and cost-effective Molecular mapping and analysis of QTL have been
carried out for a number of qualitative and quantitative traits including
stress tolerance which has undoubtedly resulted in a great magnitude
of knowledge and better understanding of the causal genetic phenom-
ena that regulate these traits However limited success has resulted by
using this knowledge to manipulate genes in an effective way for the
improvement of a speci1047297c trait in crops although some crop cultivars
developed through using these tools have performed extremely well
under1047297
eld stressconditions However there are a variety of reasons forthe limited success in terms of achieving enhanced drought tolerance
using the marker-assisted breeding approach For example a major
dif 1047297culty confronting the scientists is the challenge of precise QTL
identi1047297cation In addition a substantial genetictimes environment interac-
tion inconsistent repeatability large number of genes regulating yield
and invalid use of mapping populations have hampered the pursuits
involving mapping of QTL for enhanced drought stress tolerance
Despite these other factors also impede the application of QTL for
genetic improvement of a trait For example due to unfavorable
epistatic interaction it is hard to transfer the effects of a desired allele to
an elite background material (Podlich et al 2004 Collins et al 2008)
Furthermore in some cases QTL from a speci1047297c background do not
show signi1047297cant effects or cease completely in different backgrounds
even under analogous growth conditions (Cho and Hong 2006 Collins
et al 2008) This makes the utilization of QTL more intricate
Transformation of the knowledge acquired from QTL-oriented molec-
ular studies into a well-de1047297ned upshot for the stakeholders is one of the
key challenges confronting the breeders Certainly a multidisciplinary
approach including more speci1047297cally the identi1047297cation of QTL mediated
signal transduction in response to stresses needs to be adopted
Furthermore although QTL cloning procedures are unwieldy they can
lead to a meaningful outcome because a cloned QTL can offer a reliable
markerfor MABas well as furnish an outlinefor thedetection of superior
allelic variants in crop species (Till et al 2007)
Genetic engineering (Transgenic approach) offers a promise whereby
one can expect a substantial improvement in a desired trait within the
shortest stretch of time Engineering genes encoding organic osmolytes
plant growth regulators antioxidants late embryogenesis abundant pro-
teins and transcription factors has resulted into transgenic lines whichhave out-performed under controlled stress conditions In most cases the
claims made by the researchers in terms of performance of transgenic
lines tested under controlled conditions are considerably in1047298ated
However with the exception of a very few notable cases most of the
transgenic lines of differentcrops have been rarely1047297eld-tested Thushow
far these transgenic lines perform under 1047297eld stress conditions is not
known because under natural1047297eld conditions a linecultivar has to face a
multitude of environmental factors other than the drought stress Thus
appraisal of performance of a transgenic line under natural 1047297eld con-
ditionsis mandatory forthe stakeholdersperspective It is also imperative
to note that most of the drought tolerant transgenic lines have been
developed using a single gene transformation which may not be as
productive as if it had been developed using transformation of many
genes Thus transferring a number of prominent genes effectively in-volved in stress tolerance to transgenic plants seems to be a logical
approach Although a large number of genes appear to be involved in
stress tolerance and most of them have been fully characterized the
function of many of them in the mechanism of stress tolerance is yet to be
investigated
References
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Ashraf M Breeding for salinity tolerance in plants Crit Rev Plant Sci 19941317 ndash42Ashraf M Akram NA Improving salinity tolerance of plants through conventional
breeding and genetic engineering an analytical comparison Biotechnol Adv200927744ndash52 doi101016jbiotechadv200905026
Ashraf M Foolad MR Roles of glycinebetaine and proline in improving plant abioticstress resistance Environ Exp Bot 200759206ndash16
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Babu RC Zhang J Blum A Ho THD Wu R Nguyen HT HVA1 a LEA gene from barleyconfers dehydration tolerance in transgenic rice( Oryzasativa L)via cell membraneprotection Plant Sci 2004166855ndash62
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tobacco chloroplasts enhances the tolerance to salt stress and water de1047297cit PhysiolPlant 2004121231ndash8Badu-Apraku B Yallou CG Registration of striga-resistant and drought tolerant tropical
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Bhatnagar-Mathur P Devi MJ Vadez V Sharma KK Differential antioxidative responses intransgenic peanut bear no relationship to their superior transpiration ef 1047297ciency underdrought stress J Plant Physiol 2009166(11)1207ndash17 doi101016jjplph200901001
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Bidinger FR Nepolean T Hash CT Yadav RS Howarth CJ Identi1047297cation of QTLs for grainyield of pearl millet (Pennisetum glaucum (L) R Br) in environments with variablemoisture during grain 1047297lling Crop Sci 200747969ndash80
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binding transcription factor conferred drought and high-salt tolerance intransgenic plants Biochem Biophys Res Commun 2007353299ndash305
Cheng Z Targolli J Huang X Wu R Wheat LEA genes PMA80 and PMA1959 enhancedehydration tolerance of transgenic rice (Oryza sativa L) Mol Breed 20021071ndash82
ChoEK Hong ChB Over-expression of tobacco NtHSP70-1 contributes to drought-stresstolerance in plants Plant Cell Reports 200625349ndash58
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Gubis J Vaňkovaacute R Červenaacute V Draguacuteňovaacute M Hudcovicovaacute M Lichtnerovaacute H et alTransformedtobacco plants with increased toleranceto droughtSouthAfr J Bot200773505ndash11
GuoO Zhang J GaoQ Xing SLi F Wang WDrought tolerancethrough over-expressionof mono ubiquitin in transgenic tobacco J Plant Physiol 20081651745 ndash55
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Harris K Klein R Mullet J Sorghum stay-green QTL individually reduces post-1047298oweringdrought-induced leaf senescence J Exp Bot 200758327ndash38
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Salvi S Tuberosa R To clone or not to clone plant QTLs present and future challengesTrends Plant Sci 200510297ndash304
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Sari-Gorla M Krajewski P Di Fonzo N Villa M Frova C Genetic analysis of droughttolerance in maize by molecular markers II Plant height and 1047298owering Theor ApplGenet 199999289ndash95
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Serraj R Sinclair TR Osmolyte accumulation can it really increase crop yield underdrought conditions Plant Cell Environ 200225333ndash41
Serraj R Krishnamurthy L Kashiwagi J Kumar J Chandra S Crouch JH Variation in roottraits of chickpea (Cicer arietinum L) grown under terminal drought Field CropsRes 200488115ndash27
Serraj R Hash CT Rizvi MHS Sharma A Yadav RS Bidinger FR Recent advances in marker-assisted selection for drought tolerance in pearl millet Plant Prod Sci 20058(3)334ndash7
Shen YG Du BX Zhang WK Zhang JS Chen SY AhCMO regulated by stresses in Atriplexhortensis can improve drought tolerance in transgenic tobacco Theor Appl Genet2002105815ndash21
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Singh SP Teran H Gutierrez JA Registration of SEA 5 and SEA 13 drought tolerant drybean germplasm Crop Sci 200141276ndash7
Singh SP Teran H Lema M Schwartz HF Miklas PN Registration of white moldresistant dry bean germplasm line A 195 J Plant Registr 2007162ndash3
Sivamani E Bahieldin A Wraith JM Improved biomass productivity and water useef 1047297ciency under water de1047297cit conditions in transgenic wheat constitutivelyexpressing the barley HVA1 gene Plant Sci 20001551ndash9
Skovmand B Reynolds MP DeLacy IH Searching genetic resources for physiologicaltraits with potential for increasing yield In Reynolds MP Ortiz-Monasterio IMcNab A editors Application of Physiology in Wheat Breeding 2001 p 17ndash28
Steele K Novel upland rice variety bred using marker-assisted selection and client-oriented breeding released in Jharkhand India Bangor University 2009
Steele KA Price AH Shashidar HE Witcombe JR Marker-assistedselection to introgressrice QTLs controlling root traitsinto an Indianupland rice variety Theor Appl Genet2006112208ndash21
Steele KA Virk DS Kumar R Prasad SC Witcombe JR Field evaluation of upland ricelines selected for QTLs controlling root traits Field Crops Res 2007101180 ndash6
Sunkar R Kapoor A Zhu JK Post transcriptional induction of two CuZn superoxidedismutase genes in Arabidopsis is mediated by down regulation of miR398 andimportant for oxidative stress tolerance Plant Cell 2006182051ndash65
Talame V Sanguineti MC Chiapparino E Bahri H Ben Salem M Forster BP et alIdenti1047297cation of Hordeum spontaneum QTL alleles improving 1047297eld performance of barley grown under rainfed conditions Ann Appl Bot 2004144309ndash20
Tanner NKCordinO BanroquesJ DoereM Linder PThe Q Motif a newlyidenti1047297ed motif in DEAD box helicases may regulate ATP binding and hydrolysis Mol Cell 200311127ndash38
Taramino G Tarchini R Ferrario S Lee M Pe ME Characterization and mapping of simplesequence repeats (SSRs) in Sorghum bicolor Theor Appl Genet 19979566ndash72
Teulat B Monneveux P Wery J Borriegraves C Souyris I Charrier A et al Relationshipsbetween relative water content andgrowth parameters in barleya QTLstudy NewPhytol 199713799-107
Thi Lang N Chi Buu B Fine mapping for drought tolerance in rice ( Oryza sativa L)Omonrice 2008169-15Till BJ Comai L Henikoff S Tillering and ecotillering for crop improvement In
Varshney RK Tuberosa R editors Genomics-assisted crop improvement genomicsapproaches and platforms vol 1 2007 p 333ndash50
Tondelli A Francia E Barabaschi D Aprile A Skinner JS Stockinger EJ et al Mappingregulatory genes as candidates for cold and drought stress tolerance in barleyTheor Appl Genet 2006112445ndash54
Tuberosa R Salvi S Genomics approaches to improve drought tolerance in cropsTrends Plant Sci 200611405ndash12
Valkoun JJ Wheat pre-breeding using wild progenitors Euphytica 200111917ndash23Vashisht AA Tuteja N Stress responsive DEAD-box helicases a new pathway to
engineer plant stress tolerance J Photochem Photobiol B Biol 200684150 ndash60Vendruscolo ECG Schuster I Pileggi M Scapim CA Molinari HBC Marur CJ et al Stress-
induced synthesis of proline confers tolerance to water de1047297cit in transgenic wheat J Plant Physiol 20071641367ndash76
Vienne D Leonardi A Damerval C Zivy M Genetics of proteome variation for QTL characterization application to drought stress responses in maize J Exp Bot 199950303ndash9
Villareal RL Mujeeb-Kazi A Rajaram S Toro ED Morphological variability in somesynthetic hexaploid wheats derived from Triticum turgidum times T tauschii J GenetBreed 1994487-16
Vinh NT Paterson AH Genome mapping and its implication for stress resistance inplants In Ashraf M Harris PJC editors Abiotic stresses plant resistance throughbreeding and molecular approaches 2005
Vinocur B Altman A Recent advances in engineering plant tolerance to abiotic stressachievements and limitations Curr Opin Biotechnol 200516123ndash32
Wang YJ Hao YJ Zhang ZG Chen T Zhang JS Chen SY Isolation of trehalose-6-phosphate phosphatase gene from tobacco and its functional analysis in yeast cells
J Plant Physiol 2005162215ndash23Wu R Garg A Engineering rice plants with trehalose-producing genes improves
tolerance to drought salt and low temperature ISB News Report 2003Xiao B Huang Y Tang N Xiong L Overexpression of LEA gene in rice improves drought
resistance under 1047297eld conditions Theor Appl Genet 200711535ndash46Xinglai P Sangang X Qiannying P Yinhong S Registration of lsquo Jinmai 50rsquo wheat Crop Sci
200646983ndash5Xiong L Lee H Ishitani M Zhu JK Regulation of osmotic stress-responsive gene
expression by the LOS6ABA1 locus in Arabidopsis J Biol Chem 20022778588ndash96Xu GW Magill CW Shertz KF Hart GE A RFLP linkage map of Sorghum bicolor (L)
Moench Theor Appl Genet 199489139ndash45Xu D Duan X Wang B Hong BHo THD Wu RExpression of a late embryogenesis abundant
protein gene HVA1 from barley confers tolerance to water de1047297cit and salt stress intransgenic rice Plant Physiol 1996110249ndash57
Yadav RS Hash CT Bidinger FR Devos KM Howarth CJ Genomic regions associatedwith grain yield and aspects of post-1047298owering drought tolerance in pearl milletacross stress environments and testers background Euphytica 2004136265ndash77
YamadaM MorishitaH Urano K Shiozaki N Yamaguchi-Shinozaki K ShinozakiK et alEffects of free proline accumulation in petunias under drought stress J Exp Bot2005561975ndash81
Yang WJ Rich PJ Axtell JD Wood KV Bonham CC Ejeta G et al Genotypic variation forglycinebetaine in sorghum Crop Sci 200343162ndash9
Yan-Ying QU Ping MU Xue-Qin L Yu-Xiu T Feng W Hong-Liang Z et al QTL mappingand correlations between leaf water potential and drought resistance in rice underupland and lowland environments Acta Agron Sin 200834(2)198ndash206
Ye XDWu XLZhaoH Frehner MNoumlsberger J Potrykus Iet al Altered fructan accumulationin transgenic Lolium multi 1047298orum plants expressing a Bacillus subtilis sacB gene Plant Cell
Rep 200120205ndash
12Zaharieva M Gaulin E Havaux M Acevedo E Monneveux P Drought and heat responses in
the wild wheat relative Aegilops geniculata Roth potential interest for wheatimprovement Crop Sci 2001411321ndash9
Zhang J Zheng HGAartiA Pantuwan G NguyenTT Tripathy JNet alLocatinggenomicregions associated with components of drought resistance in rice comparativemapping within and across species Theor Appl Genet 200110319ndash29
Zhang GH Su Q An LJ Wu S Characterization and expression of a vacuolar Na+H+
antiporter gene from the monocot halophyte Aeluropus littoralis Plant PhysiolBiochem 200846117ndash26
Zhao J QTLs for oil content and their relationships to other agronomic traits in anEuropeantimesChinese oilseed rape population Germany Diss Grorg-Agust Univer-sity of Goettingen 2002
Zhao J Ren W Zhi D Wang L Xia G Arabidopsis DREB1ACBF3 bestowed transgenic tallfescue increased tolerance to drought stress Plant Cell Rep 2007261521ndash8
Zhou W Li Y Zhao BC Ge RC Shen YZ Wang G Huang ZJ Over-expression of TaSTRG gene improves salt and drought tolerance in rice J Plant Physiol 200915(166)1660ndash70
and cost-effective Molecular mapping and analysis of QTL have been
carried out for a number of qualitative and quantitative traits including
stress tolerance which has undoubtedly resulted in a great magnitude
of knowledge and better understanding of the causal genetic phenom-
ena that regulate these traits However limited success has resulted by
using this knowledge to manipulate genes in an effective way for the
improvement of a speci1047297c trait in crops although some crop cultivars
developed through using these tools have performed extremely well
under1047297
eld stressconditions However there are a variety of reasons forthe limited success in terms of achieving enhanced drought tolerance
using the marker-assisted breeding approach For example a major
dif 1047297culty confronting the scientists is the challenge of precise QTL
identi1047297cation In addition a substantial genetictimes environment interac-
tion inconsistent repeatability large number of genes regulating yield
and invalid use of mapping populations have hampered the pursuits
involving mapping of QTL for enhanced drought stress tolerance
Despite these other factors also impede the application of QTL for
genetic improvement of a trait For example due to unfavorable
epistatic interaction it is hard to transfer the effects of a desired allele to
an elite background material (Podlich et al 2004 Collins et al 2008)
Furthermore in some cases QTL from a speci1047297c background do not
show signi1047297cant effects or cease completely in different backgrounds
even under analogous growth conditions (Cho and Hong 2006 Collins
et al 2008) This makes the utilization of QTL more intricate
Transformation of the knowledge acquired from QTL-oriented molec-
ular studies into a well-de1047297ned upshot for the stakeholders is one of the
key challenges confronting the breeders Certainly a multidisciplinary
approach including more speci1047297cally the identi1047297cation of QTL mediated
signal transduction in response to stresses needs to be adopted
Furthermore although QTL cloning procedures are unwieldy they can
lead to a meaningful outcome because a cloned QTL can offer a reliable
markerfor MABas well as furnish an outlinefor thedetection of superior
allelic variants in crop species (Till et al 2007)
Genetic engineering (Transgenic approach) offers a promise whereby
one can expect a substantial improvement in a desired trait within the
shortest stretch of time Engineering genes encoding organic osmolytes
plant growth regulators antioxidants late embryogenesis abundant pro-
teins and transcription factors has resulted into transgenic lines whichhave out-performed under controlled stress conditions In most cases the
claims made by the researchers in terms of performance of transgenic
lines tested under controlled conditions are considerably in1047298ated
However with the exception of a very few notable cases most of the
transgenic lines of differentcrops have been rarely1047297eld-tested Thushow
far these transgenic lines perform under 1047297eld stress conditions is not
known because under natural1047297eld conditions a linecultivar has to face a
multitude of environmental factors other than the drought stress Thus
appraisal of performance of a transgenic line under natural 1047297eld con-
ditionsis mandatory forthe stakeholdersperspective It is also imperative
to note that most of the drought tolerant transgenic lines have been
developed using a single gene transformation which may not be as
productive as if it had been developed using transformation of many
genes Thus transferring a number of prominent genes effectively in-volved in stress tolerance to transgenic plants seems to be a logical
approach Although a large number of genes appear to be involved in
stress tolerance and most of them have been fully characterized the
function of many of them in the mechanism of stress tolerance is yet to be
investigated
References
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Ashraf M Breeding for salinity tolerance in plants Crit Rev Plant Sci 19941317 ndash42Ashraf M Akram NA Improving salinity tolerance of plants through conventional
breeding and genetic engineering an analytical comparison Biotechnol Adv200927744ndash52 doi101016jbiotechadv200905026
Ashraf M Foolad MR Roles of glycinebetaine and proline in improving plant abioticstress resistance Environ Exp Bot 200759206ndash16
Ashraf M Athar HR Harris PJC Kwon TR Some prospective strategies for improvingcrop salt tolerance Adv Agron 20089745-110
Asins MJ Present and future of quantitative trait locus analysis in plant breeding PlantBreed 2002121281ndash91
Babu RC Nguyen BD Chamarerk V Shanmugasundaram P Chezhian P Jeyaprakash P et alGeneticanalysis of droughtresistance in riceby molecular markers association betweensecondary traits and 1047297eld performance Crop Sci 2003431457ndash69
Babu RC Zhang J Blum A Ho THD Wu R Nguyen HT HVA1 a LEA gene from barleyconfers dehydration tolerance in transgenic rice( Oryzasativa L)via cell membraneprotection Plant Sci 2004166855ndash62
Badawi GH Kawano N Yamauchi Y Over-expression of ascorbate peroxidase in
tobacco chloroplasts enhances the tolerance to salt stress and water de1047297cit PhysiolPlant 2004121231ndash8Badu-Apraku B Yallou CG Registration of striga-resistant and drought tolerant tropical
early maize populations TZE-W Pop DT STR C4 and TZE-Y Pop DT STR C4 J PlantRegistr 20093(1)86ndash90
Badu-Apraku B Menkir A Kling JG Fakorede MAB Registration of 16 striga resistantearly maturing tropical maize inbred lines Crop Sci 2006461410ndash1
Baenziger PS Beecher B Graybosch RA Ibrahim AMH Baltensperger DD Nelson LA et alRegistration of lsquoNEO1643rsquo wheat J Plant Registr 20082(1)36ndash42
Baumlnziger M Setimela PS Hodson D Vivek B Breeding for improved drought tolerance inmaize adapted to southern Africa Proceedings of the 4th International Crop ScienceCongress Brisbane Australia Published on CDROM 2004 26 Sep ndash 1 Oct
Bartels D Sunkar R Drought and salt tolerance in plants Crit Rev Plant Sci 20052423 ndash58Baum M Grandol S Backes G Jahoor A Sabbagh A Ceccarelli S QTLs for agronomic
traits in the Mediterranean environment identi1047297ed in recombinant inbred lines of the cross lsquoArtarsquo H spontaneum 41-1 Theor Appl Genet 20031071215ndash25
Bergman JW Riveland NR Flynn CR Carlson GR Wichman DM Registration of lsquoMorlinrsquo
saf 1047298ower Crop Sci 2001411640Bernier J Kumar A Venuprasad R Spaner D Atlin G A large-effect QTL for
grain yield under reproductive-stage drought stress in upland rice Crop Sci200747507ndash18
BernierJ Kumar A SerrajR SpanerD Atlin G Review breeding uplandrice fordroughtresistance J Sci Food Agric 200888927ndash39
Bernier J Serraj R Kumar A Venuprasad R Impa S Gowdaa RPV et al The large-effectdrought-resistance QTL qtl121 increases water uptake in upland rice Field CropsRes 200911039ndash46
Bhatnagar-Mathur P ReddyDS Lavanya M Yamaguchi-Shinozaki K Sharma KK Stress-inducible expression of Arabidopsis thaliana DREB1A in transgenic peanut ( Arachishypogaea L) increases transpiration ef 1047297ciency under water-limiting conditionsPlant Cell Rep 2007262071ndash82
Bhatnagar-Mathur P Devi MJ Vadez V Sharma KK Differential antioxidative responses intransgenic peanut bear no relationship to their superior transpiration ef 1047297ciency underdrought stress J Plant Physiol 2009166(11)1207ndash17 doi101016jjplph200901001
Bidinger FR Serraj R Rizvi SMH Howarth C Yadav RS Hash CT Field evaluation of droughttolerance QTL effects on phenotype and adaptation in pearl millet [ Pennisetum glaucum(L) R Br] topcross hybrids Field Crops Res 200594(1)14ndash32
Bidinger FR Nepolean T Hash CT Yadav RS Howarth CJ Identi1047297cation of QTLs for grainyield of pearl millet (Pennisetum glaucum (L) R Br) in environments with variablemoisture during grain 1047297lling Crop Sci 200747969ndash80
Bowers JE Abbey C Anderson S Chang C Draye X A high-density geneticrecombination map of sequence-tagged sites for Sorghum as a framework forcomparative structural and evolutionary genomics of tropical grains and grassesGenetics 2003165367ndash86
Brick MA Ogg JB Singh SP Schwartz HF Johnson JJ Pastor-Corrales MA Registration of drought-tolerant rust-resistant high-yielding pinto bean germplasm lineCO46348 J Plant Registr 20082(2)120ndash4
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Busk PK Pages M Regulation of abscisic acid induced transcription Plant Mol Biol199837425ndash35
Carena MJ Wanner DW Development of genetically broad-based inbred lines of maizefor early-maturing (70-80RM) hybrids J Plant Registr 20093107ndash11
Cash SDBrucknerPL Wichman DMKephart KD Berg JEBoynerR et alRegistration of Willow Creek forage wheat J Plant Registr 20093(2)185ndash90
Cattivelli L Rizza F Badeck FW Mazzucotelli E Francia AMEM Mare AT et al Droughttolerance improvement in crop plants an integrated view from breeding to genomics
Field Crops Res 20081051-14Chen M Wang QY Cheng XG Xu ZS Li LC Ye XG et al GmDREB2 a soybean DRE-
binding transcription factor conferred drought and high-salt tolerance intransgenic plants Biochem Biophys Res Commun 2007353299ndash305
Cheng Z Targolli J Huang X Wu R Wheat LEA genes PMA80 and PMA1959 enhancedehydration tolerance of transgenic rice (Oryza sativa L) Mol Breed 20021071ndash82
ChoEK Hong ChB Over-expression of tobacco NtHSP70-1 contributes to drought-stresstolerance in plants Plant Cell Reports 200625349ndash58
Close TJ Dehydrins a commonality in the response of plants to dehydration and lowtemperature Physiol Plant 1997100291ndash6
Collins NC Tardieu F Tuberosa R Quantitative trait loci and crop performance underabiotic stress where do we stand Plant Physiol 2008147469ndash86
Concept Note Combining breeding and biotechnology to develop water ef 1047297cient maizefor Africa (WEMA) Afr Agric Technol Foundation
Courtois BShen L Petalcorin WCarandang S MauleonR LiZ Locating QTLs controllingconstitutive root traits in the rice population IAC 165-Co39 Euphytica 2003134335ndash45
Da-hong L Hui L Yang YL Ping-ping Z Jian-sheng L Down-regulated expression of RACK1geneby RNAinterference enhancesdroughttolerancein rice RiceSci 200916(1)14ndash20
Dalal M Tayal D Chinnusamy V Bansala KC Abiotic stress and ABA-inducible group 4 LEAfrom Brassicanapus plays a key rolein saltand droughttolerance J Biotechnol 2009139137ndash45
Ding Z Li S An X Liu X Qin H Wang D Transgenic expression of MYB15 confers enhancedsensitivity to abscisic acid and improved drought tolerance in Arabidopsis thaliana
J Genet Genomics 20093617ndash29Dubouzet JG Sakuma Y Ito Y Kasuga M Dubouzet EG Miura S et al OsDREB genes in
rice Oryza sativa L encode transcription activators that function in drought high-salt- and cold-responsive gene expression Plant J 200333751ndash63
El-Far IA Allan AY Responses of some wheat cultivars to sowing methods and droughtat different stages of growth Assuit J Agric Sci 199526(1)267ndash77
Eltayeb AE Kawano N Badawi GH Kaminaka H Sanekata T Shibahara T et alOverexpression of monodehydroascorbate reductase in transgenic tobacco confersenhanced tolerance to ozonesalt and polyethyleneglycol stresses Planta 2007225(5)1255ndash64
Falconer DS Introduction to quantitative genetics London New York Longman 1989Feng-ling FU Zhi-Lei F Shi-bing G Shu-feng Z Wan-chen L Evaluation and quantitative
inheritanceof several drought-relativetraits in maize Agric SciChina 20087(3)280ndash90Finkelstein R Gampala S Rock C Abscisic acid signaling in seeds and seedlings Plant
Cell 20021415ndash45Flowers TJ Improving crop salt tolerance J Exp Bot 200455307ndash19Fujita Y Fujita M Satoh R Maruyama K Parvez MM Seki M et al AREB1 is a transcription
Giraudat J Parcy F Bertauche N Gosti F Leung J Morris PC et al Current advances inabscisic acid action and signalling Plant Mol Biol 1994261557ndash77
Gorbalenya AE Koonin EV Helicases amino acid sequence comparisons and structurendashfunction relationships Curr Opin Struct Biol 19933419ndash29
Gosal SS Wani SH Kang MS Biotechnology and drought tolerance J Crop Improvement20092319ndash54
Gubis J Vaňkovaacute R Červenaacute V Draguacuteňovaacute M Hudcovicovaacute M Lichtnerovaacute H et alTransformedtobacco plants with increased toleranceto droughtSouthAfr J Bot200773505ndash11
GuoO Zhang J GaoQ Xing SLi F Wang WDrought tolerancethrough over-expressionof mono ubiquitin in transgenic tobacco J Plant Physiol 20081651745 ndash55
Haley SD Johnson JJ Peairs FB Quick JS Stromberger JA Clayshulte SR et al Registration of lsquoRipperrsquo wheat J Plant Registr 200711ndash6
Han SE Park SR Kwon HB Yi BY Lee GB Byun MO Genetic engineering of drought-resistant tobacco plants by introducingthe trehalose phosphorylase (TP) gene fromPleurotus sajor-caju Plant Cell Tissue Organ Cult 200582151ndash8
Harris K Klein R Mullet J Sorghum stay-green QTL individually reduces post-1047298oweringdrought-induced leaf senescence J Exp Bot 200758327ndash38
Hong BS Zong-Suo L Ming-An S LEA proteins in higher plants structure functiongene expression and regulation Colloids Surf B Biointerf 200545131ndash5
Howarth CJ Yadav RS Successful marker assisted selection for drought tolerance anddisease resistance in pearl milletIGER Innovations 2002
Humphreys MO Humphreys MW Breeding for stress resistance general principles InAshraf M Harris PJC editors Abiotic stresses plant resistance through breedingand molecular approaches 2005 p 19ndash46
Ingram J Bartels D The molecular basis of dehydration tolerance in plants Ann RevPlant Physiol Plant Mol Biol 199647377ndash403
Jaglo-Ottosen KR Gilmour SJ Zarka DG Schabenberger O Thomashow MF ArabidopsisCBF1 overexpression induces COR genes and enhances freezing tolerance Science1998280104ndash6
Jang IC Oh SJ Seo JS Choi WB Song SI Kim CH et al Expression of a bifunctional fusion of the Escherichia coli genes for trehalose-6-phosphate synthase and trehalose-6-phosphate phosphatase in transgenic rice plants increases trehalose accumulationandabiotic stresstolerancewithoutstuntinggrowth Plant Physiol2003131516ndash24
Jeanneau M Gerentes D Foueillassar X Zivy M Vidal J Toppan A et al Improvement of drought tolerance in maize towards the functional validation of the Zm-Asr1 gene andincrease of water use ef 1047297ciency by over-expressing C4-PEPC Biochimie 2002841127ndash35
Jenson KB Palazzo AJ Waldron BL Bushman BS Registration of lsquoFirstStrikersquo Slenderwheatgrass J Plant Registr 2007124ndash5
Joung-youn K Hyung-in C Min-young I Soo-young K Arabidopsis basic leucine zipperproteins that mediate stress-responsive abscisic acid signaling Plant Cell 20024343ndash57
Juskiw PE Helm JH Oro M Nyachiro JM Salmon DF Registration of lsquoBentleyrsquo barley
J Plant Registr 20093(2)119ndash23Kalamaki MS Alexandrou D Lazari D Merkouropoulos G Fotopoulos V Pateraki I et al
Over-expression of a tomato N-acetyl-L -glutamate synthase gene (SlNAGS1) in Arabidopsis thaliana results in high ornithine levels and increased tolerance in saltand drought stresses J Exp Bot 200960(6)1859ndash71
Kamoshita A Babu CR Boopathi NM Fukai S Phenotypic and genotypic analysis of drought-resistance traits for development of rice cultivars adapted to rainfedenvironments Field Crops Res 2008109(103)1-23 doi101016jfcr200806010
KarakasB Ozias-AkinsP Stushnoff C SuefferheldM Rieger M Salinityand drought toleranceof mannitol-accumulating transgenic tobacco Plant Cell Environ 199720609ndash16
Karim S Aronsson H Ericson H Pirhonen M Leyman B Welin B et al Improved droughttolerance without undesired side effects in transgenic plants producing trehalose PlantMol Biol 200764371ndash86
Kasuga M Liu Q Miura S Yamaguchi-Shinozaki K Shinozaki K Improving plant droughtsaltand freezing tolerance by gene transfer of a single stress-inducible transcription factorNat Biotechnol 199917287ndash91
Kindiger M Gaub H Hasegawac M Katsurab Y Ueyamad K Gotob S et al Registrationof lsquoNanryorsquo tall fescue Crop Sci 2006461815ndash6
Kong I Dong J HartGE Characteristics linkage mappositionsand allelic differentiationof Sorghum bicolour (L) Moench by DNA simple-sequence repeats (SSRs) TheorAppl Genet 2000101438ndash48
Kumar R Venuprasad R Atlin GN Genetic analysis of rainfed lowland rice droughttolerance under naturally-occurring stress in eastern India heritability and QTL effects Field Crops Res 200710342ndash52
La1047297tte HR Price AH Courtois B Yield response to water de1047297cit in an upland ricemapping population associations among traits and genetic markers Theor ApplGenet 20041091237ndash46
Lal S Gulyani V Khurana P Overexpressionof HVA1 gene frombarley generates tolerance tosalinity and water stress in transgenic mulberry (Morus indica) Transgenic Res 200817
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63Lanceras J Pantuwan G Jongdee B Toojinda T Quantitative trait loci associated withdrought tolerance at reproductive stage in rice Plant Physiol 2004135384ndash99
Levi A Ovnat L Paterson AH Saranga Y Photosynthesis of cotton near-isogenic linesintrogressed with QTLs for productivity and drought related traits Plant Sci 2009a17788ndash96
Levi A Paterson AH Barak V Yakir D Wang B Chee PW et al Field evaluation of cottonnear-isogenic lines introgressed with QTLs for productivity and drought relatedtraits Mol Breed 2009b23179ndash95
Liu Q Kasuga M Sakuma Y Abe H Miura S Yamaguchi- Shinozaki K et al Twotranscription factors DREB1 and DREB2 with an EREBPAP2 DNA binding domainseparate two cellular signal transduction pathways in drought-and low-temper-ature-responsive gene expression respectively in Arabidopsis Plant Cell 1998101391ndash406
Liu X Hua X Guo J Qi D Wang L Liu Z et al Enhanced tolerance to drought stress intransgenic tobacco plants overexpressing Biotechnol Lett 2008301275ndash80
Liu X Wanga Z Wanga L Wua R Phillips J Deng X LEA 4 group genes from theresurrection plant Boea hygrometrica confer dehydration tolerance in transgenictobacco Plant Sci 200917690ndash8
Luchi S Kobayashi M Taji T Naramoto M Seki M Kato T et al Regulation of droughttolerance by gene manipulation of 9-cis-epoxycarotenoid dioxygenase a keyenzyme in abscisic acid biosynthesis in Arabidopsis Plant J 200127325ndash33
Luo Y Liu YB Dong YX Gao XQ Zhang YS Expression of a putative alfalfa helicase increasestolerance to abioticstress in Arabidopsis by enhancing the capacities for ROS scavengingand osmotic adjustment J Plant Physiol 2009166385ndash94
MacLean JLDaweDC Hardy B Hettel GPRiceAlmanac sourcebook for themostimportanteconomic activity on Earth3rd ed Wallingford England CABI Publishing 2002
Mansour MMF Nitrogen containing compounds and adaptation of plants to salinitystress Biol Plant 200043491ndash500
McKersie BD Bowley SR Harjanto E Leprince O Water-de1047297cit tolerance and 1047297eldperformance of transgenic alfalfa overexpressing superoxide dismutase Plant Physiol19961111177ndash81
McKersie BD Murnaghan J Bowley SR Manipulating freezing tolerance in transgenicplants Acta Physiol Plant 199719485ndash95
Miranda JA Avonce N Suaacuterez R Thevelein JM Dijck PV Iturriaga G A bifunctional TPS-TPP enzyme from yeast confers tolerance to multiple and extreme abiotic-stressconditions in transgenic Arabidopsis Planta 2007226(6)1411ndash21
Mohammadi M Taleei A Zeinali H Naghavi MR Ceccarelli S Grando Baum M QTL analysis for phenologic traits in doubled haploid population of barley Int J AgricBiol 20057(5)820ndash3
Mohanty A Kathuria H Ferjani A Sakamoto A Mohanty P Murata N et al Transgenicsof an elite indica rice variety Pusa Basmati 1 harbouring the codA gene are highlytolerant to salt stress Theor Appl Genet 200210651ndash7
Noaman MM El Sayad AA Asaad FA El Sherbini AM El Bawab AO El Moselhi MA et alRegistration of lsquoGiza 126rsquo barley Crop Sci 199535(6)1710
Noaman MM Ahmed IA El-Sayed AA Abo-El-Enin RA El-Gamal AS El-Sherbiny AM et alRegistration of lsquoGiza 2000rsquo drought-tolerant six-rowed barley for rainfed and newreclaimed areas in Egypt Crop Sci 200747440
Obert DE Evans CP Wesenberg DM Windes JM Erickson CA Jackson EW et alRegistration of lsquoLenetahrsquo spring barley J Plant Registr 20082(2)85ndash7
Oh SJ Jeong JS Kim EH Yi NR Yi SI Jang IC et al Matrix attachment region fromthe chicken lysozyme locus reduces variability in transgene expression andconfers copy number-dependence in transgenic rice plants Plant Cell Rep20054145ndash54
Oh SJ Kim YS Kwon C Park HK Jeong JS Kim JK Overexpression of the transcriptionfactor AP37 in rice improves grain yield under drought conditions Plant Physiol
20092191ndash
200 doi101104pp109137554Owttrim GW RNA helicases and abiotic stress Nucleic AcidsRes 200634(11)3220ndash30Park BJ Liu Z Kanno A Kameya T Increased tolerance to salt and water de1047297cit stress in
transgenic lettuce (Lactuca sativa L) by constitutive expression of LEA PlantGrowth Regul 2005a45165ndash71
Park BJ Liu Z Kanno A Kameya T Genetic improvement of Chinese cabbage for salt anddroughttolerance by constitutiveexpression of a B napus LEAgenePlantSci 2005b169553ndash8
Pastori GM Foyer CH Common componentsnetworks and pathways of crosstoleranceto stress The central role of ldquoredoxrdquo and abscisic acid-mediated controls PlantPhysiol 2002129460ndash8
PellegrineschiA ReynoldsM PachecoM Brito RMAlmeraya R Yamaguchi-ShinozakiK et alStress-induced expression in wheat of the Arabidopsis thaliana DREB1Agenedelayswaterstress symptoms under greenhouse conditions Genome 200447493ndash500
Perl A Perl-Treves R Galili S Aviv D Shalgi E Malkin S et al Enhanced oxidative-stressdefense in transgenic potato overexpressing tomato Cu Zn superoxide dismutaseTheor Appl Genet 199385568ndash76
Podlich DW Winkler CR Cooper M Mapping as you go an effective approach formarker-assisted selection of complex traits Crop Sci 2004441560ndash71
182 M Ashraf Biotechnology Advances 28 (2010) 169ndash183
Pratt RC Casey MA Registration of maize germplasm line Oh605 Crop Sci 2006461004ndash5Qin F Kakimoto M Sakuma Y Maruyama K Osakabe Y Tran LSP et al Regulation and
functional analysis of ZmDREB2A in response to drought and heat stress in Zea mays LPlant J 20075054ndash69
Quan R Shang M Zhang H Zhao Y Zhang J Engineering of enhanced glycinebetainesynthesis improves drought tolerance in maize Plant Biotechnol J 20042477ndash86
Quarrie SA Gulli M Calestani C Steed A Marmiroli N Location of a gene regulatingdrought-induced abscisic acid production on the long arm of chromosome 5A of wheat Theor Appl Genet 199489794ndash800
QuickJS Stromberger JA Clayshulte S Clifford B Johnson JJ Peairs FB et al Registrationof lsquoPrairie Redrsquo wheat Crop Sci 2001411362ndash3
RajaramS Prospects andpromise ofwheatbreedingin the21stcentury Euphytica 20011193-15Rajaram S Role of conventional plant breeding and biotechnology in future wheat
production Turk J Agric Forest 200529105ndash11Rami JF Dufour P Trouche G Fliedel G Mestres C Davrieux F et al Quantitative trait
loci for grain quality productivity morphological and agronomical traits insorghum (Sorghum bicolor L Moench) Theor Appl Genet 199897605ndash16
Rassaa N Salahb HBH Latiri K Thermal responses of durum wheat Triticum durum toearly water stress consequence on leaf and 1047298ower development Plant Biol Pathol2008331(5)363ndash71
Reddy LJ Nigam SN Rao RCN Reddy NS Registration of ICGV 87354 peanut germplasmwith drought tolerance and rust resistance Crop Sci 200141274ndash5
Ribaut JM Ragot M Marker-assisted selection to improve drought adaptation in maize thebackcross approach perspectives limitations and alternatives J Exp Bot 200658351ndash60
Robin S Pathan MS Courtois B La1047297tte R Carandang S Lanceras S et al Mappingosmotic adjustment in an advanced back-cross inbred population of rice Theor
Appl Genet 2003107(7)1288ndash96Romero C Belles JM Vaya JL Serrano R Culianez-Macia FA Expression of the yeast
trehalose-6-phosphate synthase gene in transgenic tobacco plants pleiotropicphenotypes include drought tolerance Planta 1997201293ndash7
Ronde JAD Cress WA Krugerd GHJ Strasserd RJ Van Staden J Photosynthetic responseof transgenic soybean plants containing an Arabidopsis P5CR gene during heat anddrought stress J Plant Physiol 20041611211ndash24
Salvi S Tuberosa R To clone or not to clone plant QTLs present and future challengesTrends Plant Sci 200510297ndash304
Sanchez AC Subudhi PK Rosenow DT Nguyen HT Mapping QTLs associated with droughtresistance in sorghum (Sorghum bicolor L Moench) Plant Mol Biol 200248713ndash26
Saranga Y Menz M Jiang CX Wright RJ Yakir D Paterson AH Genomic dissection of genotypetimesenvironment interactions conferring adaptation of cotton to aridconditions Genome Res 2001111988ndash95
Sari-Gorla M Krajewski P Di Fonzo N Villa M Frova C Genetic analysis of droughttolerance in maize by molecular markers II Plant height and 1047298owering Theor ApplGenet 199999289ndash95
Seki M Narusaka M Ishida J Nanjo T Fujita M Oono Y et al Monitoring the expressionpro1047297les of 7000 Arabidopsis genes under drought cold and high-salinity stressesusing a full-length cDNA microarray Plant J 200231279ndash92
Serraj R Sinclair TR Osmolyte accumulation can it really increase crop yield underdrought conditions Plant Cell Environ 200225333ndash41
Serraj R Krishnamurthy L Kashiwagi J Kumar J Chandra S Crouch JH Variation in roottraits of chickpea (Cicer arietinum L) grown under terminal drought Field CropsRes 200488115ndash27
Serraj R Hash CT Rizvi MHS Sharma A Yadav RS Bidinger FR Recent advances in marker-assisted selection for drought tolerance in pearl millet Plant Prod Sci 20058(3)334ndash7
Shen YG Du BX Zhang WK Zhang JS Chen SY AhCMO regulated by stresses in Atriplexhortensis can improve drought tolerance in transgenic tobacco Theor Appl Genet2002105815ndash21
Shinozaki K Yamaguchi-Shinozaki K Molecular responses to dehydration and lowtemperature differences and cross-talk between two stress signaling pathwaysCurr Opin Plant Biol 20003217ndash23
Singh KB Omar M Saxena MC Johansen C Registration of FLIP 87-59C a drought-tolerant chickpea germplasm line Crop Sci 199636(2)1ndash2
Singh SP Teran H Gutierrez JA Registration of SEA 5 and SEA 13 drought tolerant drybean germplasm Crop Sci 200141276ndash7
Singh SP Teran H Lema M Schwartz HF Miklas PN Registration of white moldresistant dry bean germplasm line A 195 J Plant Registr 2007162ndash3
Sivamani E Bahieldin A Wraith JM Improved biomass productivity and water useef 1047297ciency under water de1047297cit conditions in transgenic wheat constitutivelyexpressing the barley HVA1 gene Plant Sci 20001551ndash9
Skovmand B Reynolds MP DeLacy IH Searching genetic resources for physiologicaltraits with potential for increasing yield In Reynolds MP Ortiz-Monasterio IMcNab A editors Application of Physiology in Wheat Breeding 2001 p 17ndash28
Steele K Novel upland rice variety bred using marker-assisted selection and client-oriented breeding released in Jharkhand India Bangor University 2009
Steele KA Price AH Shashidar HE Witcombe JR Marker-assistedselection to introgressrice QTLs controlling root traitsinto an Indianupland rice variety Theor Appl Genet2006112208ndash21
Steele KA Virk DS Kumar R Prasad SC Witcombe JR Field evaluation of upland ricelines selected for QTLs controlling root traits Field Crops Res 2007101180 ndash6
Sunkar R Kapoor A Zhu JK Post transcriptional induction of two CuZn superoxidedismutase genes in Arabidopsis is mediated by down regulation of miR398 andimportant for oxidative stress tolerance Plant Cell 2006182051ndash65
Talame V Sanguineti MC Chiapparino E Bahri H Ben Salem M Forster BP et alIdenti1047297cation of Hordeum spontaneum QTL alleles improving 1047297eld performance of barley grown under rainfed conditions Ann Appl Bot 2004144309ndash20
Tanner NKCordinO BanroquesJ DoereM Linder PThe Q Motif a newlyidenti1047297ed motif in DEAD box helicases may regulate ATP binding and hydrolysis Mol Cell 200311127ndash38
Taramino G Tarchini R Ferrario S Lee M Pe ME Characterization and mapping of simplesequence repeats (SSRs) in Sorghum bicolor Theor Appl Genet 19979566ndash72
Teulat B Monneveux P Wery J Borriegraves C Souyris I Charrier A et al Relationshipsbetween relative water content andgrowth parameters in barleya QTLstudy NewPhytol 199713799-107
Thi Lang N Chi Buu B Fine mapping for drought tolerance in rice ( Oryza sativa L)Omonrice 2008169-15Till BJ Comai L Henikoff S Tillering and ecotillering for crop improvement In
Varshney RK Tuberosa R editors Genomics-assisted crop improvement genomicsapproaches and platforms vol 1 2007 p 333ndash50
Tondelli A Francia E Barabaschi D Aprile A Skinner JS Stockinger EJ et al Mappingregulatory genes as candidates for cold and drought stress tolerance in barleyTheor Appl Genet 2006112445ndash54
Tuberosa R Salvi S Genomics approaches to improve drought tolerance in cropsTrends Plant Sci 200611405ndash12
Valkoun JJ Wheat pre-breeding using wild progenitors Euphytica 200111917ndash23Vashisht AA Tuteja N Stress responsive DEAD-box helicases a new pathway to
engineer plant stress tolerance J Photochem Photobiol B Biol 200684150 ndash60Vendruscolo ECG Schuster I Pileggi M Scapim CA Molinari HBC Marur CJ et al Stress-
induced synthesis of proline confers tolerance to water de1047297cit in transgenic wheat J Plant Physiol 20071641367ndash76
Vienne D Leonardi A Damerval C Zivy M Genetics of proteome variation for QTL characterization application to drought stress responses in maize J Exp Bot 199950303ndash9
Villareal RL Mujeeb-Kazi A Rajaram S Toro ED Morphological variability in somesynthetic hexaploid wheats derived from Triticum turgidum times T tauschii J GenetBreed 1994487-16
Vinh NT Paterson AH Genome mapping and its implication for stress resistance inplants In Ashraf M Harris PJC editors Abiotic stresses plant resistance throughbreeding and molecular approaches 2005
Vinocur B Altman A Recent advances in engineering plant tolerance to abiotic stressachievements and limitations Curr Opin Biotechnol 200516123ndash32
Wang YJ Hao YJ Zhang ZG Chen T Zhang JS Chen SY Isolation of trehalose-6-phosphate phosphatase gene from tobacco and its functional analysis in yeast cells
J Plant Physiol 2005162215ndash23Wu R Garg A Engineering rice plants with trehalose-producing genes improves
tolerance to drought salt and low temperature ISB News Report 2003Xiao B Huang Y Tang N Xiong L Overexpression of LEA gene in rice improves drought
resistance under 1047297eld conditions Theor Appl Genet 200711535ndash46Xinglai P Sangang X Qiannying P Yinhong S Registration of lsquo Jinmai 50rsquo wheat Crop Sci
200646983ndash5Xiong L Lee H Ishitani M Zhu JK Regulation of osmotic stress-responsive gene
expression by the LOS6ABA1 locus in Arabidopsis J Biol Chem 20022778588ndash96Xu GW Magill CW Shertz KF Hart GE A RFLP linkage map of Sorghum bicolor (L)
Moench Theor Appl Genet 199489139ndash45Xu D Duan X Wang B Hong BHo THD Wu RExpression of a late embryogenesis abundant
protein gene HVA1 from barley confers tolerance to water de1047297cit and salt stress intransgenic rice Plant Physiol 1996110249ndash57
Yadav RS Hash CT Bidinger FR Devos KM Howarth CJ Genomic regions associatedwith grain yield and aspects of post-1047298owering drought tolerance in pearl milletacross stress environments and testers background Euphytica 2004136265ndash77
YamadaM MorishitaH Urano K Shiozaki N Yamaguchi-Shinozaki K ShinozakiK et alEffects of free proline accumulation in petunias under drought stress J Exp Bot2005561975ndash81
Yang WJ Rich PJ Axtell JD Wood KV Bonham CC Ejeta G et al Genotypic variation forglycinebetaine in sorghum Crop Sci 200343162ndash9
Yan-Ying QU Ping MU Xue-Qin L Yu-Xiu T Feng W Hong-Liang Z et al QTL mappingand correlations between leaf water potential and drought resistance in rice underupland and lowland environments Acta Agron Sin 200834(2)198ndash206
Ye XDWu XLZhaoH Frehner MNoumlsberger J Potrykus Iet al Altered fructan accumulationin transgenic Lolium multi 1047298orum plants expressing a Bacillus subtilis sacB gene Plant Cell
Rep 200120205ndash
12Zaharieva M Gaulin E Havaux M Acevedo E Monneveux P Drought and heat responses in
the wild wheat relative Aegilops geniculata Roth potential interest for wheatimprovement Crop Sci 2001411321ndash9
Zhang J Zheng HGAartiA Pantuwan G NguyenTT Tripathy JNet alLocatinggenomicregions associated with components of drought resistance in rice comparativemapping within and across species Theor Appl Genet 200110319ndash29
Zhang GH Su Q An LJ Wu S Characterization and expression of a vacuolar Na+H+
antiporter gene from the monocot halophyte Aeluropus littoralis Plant PhysiolBiochem 200846117ndash26
Zhao J QTLs for oil content and their relationships to other agronomic traits in anEuropeantimesChinese oilseed rape population Germany Diss Grorg-Agust Univer-sity of Goettingen 2002
Zhao J Ren W Zhi D Wang L Xia G Arabidopsis DREB1ACBF3 bestowed transgenic tallfescue increased tolerance to drought stress Plant Cell Rep 2007261521ndash8
Zhou W Li Y Zhao BC Ge RC Shen YZ Wang G Huang ZJ Over-expression of TaSTRG gene improves salt and drought tolerance in rice J Plant Physiol 200915(166)1660ndash70
and cost-effective Molecular mapping and analysis of QTL have been
carried out for a number of qualitative and quantitative traits including
stress tolerance which has undoubtedly resulted in a great magnitude
of knowledge and better understanding of the causal genetic phenom-
ena that regulate these traits However limited success has resulted by
using this knowledge to manipulate genes in an effective way for the
improvement of a speci1047297c trait in crops although some crop cultivars
developed through using these tools have performed extremely well
under1047297
eld stressconditions However there are a variety of reasons forthe limited success in terms of achieving enhanced drought tolerance
using the marker-assisted breeding approach For example a major
dif 1047297culty confronting the scientists is the challenge of precise QTL
identi1047297cation In addition a substantial genetictimes environment interac-
tion inconsistent repeatability large number of genes regulating yield
and invalid use of mapping populations have hampered the pursuits
involving mapping of QTL for enhanced drought stress tolerance
Despite these other factors also impede the application of QTL for
genetic improvement of a trait For example due to unfavorable
epistatic interaction it is hard to transfer the effects of a desired allele to
an elite background material (Podlich et al 2004 Collins et al 2008)
Furthermore in some cases QTL from a speci1047297c background do not
show signi1047297cant effects or cease completely in different backgrounds
even under analogous growth conditions (Cho and Hong 2006 Collins
et al 2008) This makes the utilization of QTL more intricate
Transformation of the knowledge acquired from QTL-oriented molec-
ular studies into a well-de1047297ned upshot for the stakeholders is one of the
key challenges confronting the breeders Certainly a multidisciplinary
approach including more speci1047297cally the identi1047297cation of QTL mediated
signal transduction in response to stresses needs to be adopted
Furthermore although QTL cloning procedures are unwieldy they can
lead to a meaningful outcome because a cloned QTL can offer a reliable
markerfor MABas well as furnish an outlinefor thedetection of superior
allelic variants in crop species (Till et al 2007)
Genetic engineering (Transgenic approach) offers a promise whereby
one can expect a substantial improvement in a desired trait within the
shortest stretch of time Engineering genes encoding organic osmolytes
plant growth regulators antioxidants late embryogenesis abundant pro-
teins and transcription factors has resulted into transgenic lines whichhave out-performed under controlled stress conditions In most cases the
claims made by the researchers in terms of performance of transgenic
lines tested under controlled conditions are considerably in1047298ated
However with the exception of a very few notable cases most of the
transgenic lines of differentcrops have been rarely1047297eld-tested Thushow
far these transgenic lines perform under 1047297eld stress conditions is not
known because under natural1047297eld conditions a linecultivar has to face a
multitude of environmental factors other than the drought stress Thus
appraisal of performance of a transgenic line under natural 1047297eld con-
ditionsis mandatory forthe stakeholdersperspective It is also imperative
to note that most of the drought tolerant transgenic lines have been
developed using a single gene transformation which may not be as
productive as if it had been developed using transformation of many
genes Thus transferring a number of prominent genes effectively in-volved in stress tolerance to transgenic plants seems to be a logical
approach Although a large number of genes appear to be involved in
stress tolerance and most of them have been fully characterized the
function of many of them in the mechanism of stress tolerance is yet to be
investigated
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Ashraf M Breeding for salinity tolerance in plants Crit Rev Plant Sci 19941317 ndash42Ashraf M Akram NA Improving salinity tolerance of plants through conventional
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Ashraf M Foolad MR Roles of glycinebetaine and proline in improving plant abioticstress resistance Environ Exp Bot 200759206ndash16
Ashraf M Athar HR Harris PJC Kwon TR Some prospective strategies for improvingcrop salt tolerance Adv Agron 20089745-110
Asins MJ Present and future of quantitative trait locus analysis in plant breeding PlantBreed 2002121281ndash91
Babu RC Nguyen BD Chamarerk V Shanmugasundaram P Chezhian P Jeyaprakash P et alGeneticanalysis of droughtresistance in riceby molecular markers association betweensecondary traits and 1047297eld performance Crop Sci 2003431457ndash69
Babu RC Zhang J Blum A Ho THD Wu R Nguyen HT HVA1 a LEA gene from barleyconfers dehydration tolerance in transgenic rice( Oryzasativa L)via cell membraneprotection Plant Sci 2004166855ndash62
Badawi GH Kawano N Yamauchi Y Over-expression of ascorbate peroxidase in
tobacco chloroplasts enhances the tolerance to salt stress and water de1047297cit PhysiolPlant 2004121231ndash8Badu-Apraku B Yallou CG Registration of striga-resistant and drought tolerant tropical
early maize populations TZE-W Pop DT STR C4 and TZE-Y Pop DT STR C4 J PlantRegistr 20093(1)86ndash90
Badu-Apraku B Menkir A Kling JG Fakorede MAB Registration of 16 striga resistantearly maturing tropical maize inbred lines Crop Sci 2006461410ndash1
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Baumlnziger M Setimela PS Hodson D Vivek B Breeding for improved drought tolerance inmaize adapted to southern Africa Proceedings of the 4th International Crop ScienceCongress Brisbane Australia Published on CDROM 2004 26 Sep ndash 1 Oct
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traits in the Mediterranean environment identi1047297ed in recombinant inbred lines of the cross lsquoArtarsquo H spontaneum 41-1 Theor Appl Genet 20031071215ndash25
Bergman JW Riveland NR Flynn CR Carlson GR Wichman DM Registration of lsquoMorlinrsquo
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grain yield under reproductive-stage drought stress in upland rice Crop Sci200747507ndash18
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Bhatnagar-Mathur P ReddyDS Lavanya M Yamaguchi-Shinozaki K Sharma KK Stress-inducible expression of Arabidopsis thaliana DREB1A in transgenic peanut ( Arachishypogaea L) increases transpiration ef 1047297ciency under water-limiting conditionsPlant Cell Rep 2007262071ndash82
Bhatnagar-Mathur P Devi MJ Vadez V Sharma KK Differential antioxidative responses intransgenic peanut bear no relationship to their superior transpiration ef 1047297ciency underdrought stress J Plant Physiol 2009166(11)1207ndash17 doi101016jjplph200901001
Bidinger FR Serraj R Rizvi SMH Howarth C Yadav RS Hash CT Field evaluation of droughttolerance QTL effects on phenotype and adaptation in pearl millet [ Pennisetum glaucum(L) R Br] topcross hybrids Field Crops Res 200594(1)14ndash32
Bidinger FR Nepolean T Hash CT Yadav RS Howarth CJ Identi1047297cation of QTLs for grainyield of pearl millet (Pennisetum glaucum (L) R Br) in environments with variablemoisture during grain 1047297lling Crop Sci 200747969ndash80
Bowers JE Abbey C Anderson S Chang C Draye X A high-density geneticrecombination map of sequence-tagged sites for Sorghum as a framework forcomparative structural and evolutionary genomics of tropical grains and grassesGenetics 2003165367ndash86
Brick MA Ogg JB Singh SP Schwartz HF Johnson JJ Pastor-Corrales MA Registration of drought-tolerant rust-resistant high-yielding pinto bean germplasm lineCO46348 J Plant Registr 20082(2)120ndash4
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Busk PK Pages M Regulation of abscisic acid induced transcription Plant Mol Biol199837425ndash35
Carena MJ Wanner DW Development of genetically broad-based inbred lines of maizefor early-maturing (70-80RM) hybrids J Plant Registr 20093107ndash11
Cash SDBrucknerPL Wichman DMKephart KD Berg JEBoynerR et alRegistration of Willow Creek forage wheat J Plant Registr 20093(2)185ndash90
Cattivelli L Rizza F Badeck FW Mazzucotelli E Francia AMEM Mare AT et al Droughttolerance improvement in crop plants an integrated view from breeding to genomics
Field Crops Res 20081051-14Chen M Wang QY Cheng XG Xu ZS Li LC Ye XG et al GmDREB2 a soybean DRE-
binding transcription factor conferred drought and high-salt tolerance intransgenic plants Biochem Biophys Res Commun 2007353299ndash305
Cheng Z Targolli J Huang X Wu R Wheat LEA genes PMA80 and PMA1959 enhancedehydration tolerance of transgenic rice (Oryza sativa L) Mol Breed 20021071ndash82
ChoEK Hong ChB Over-expression of tobacco NtHSP70-1 contributes to drought-stresstolerance in plants Plant Cell Reports 200625349ndash58
Close TJ Dehydrins a commonality in the response of plants to dehydration and lowtemperature Physiol Plant 1997100291ndash6
Collins NC Tardieu F Tuberosa R Quantitative trait loci and crop performance underabiotic stress where do we stand Plant Physiol 2008147469ndash86
Concept Note Combining breeding and biotechnology to develop water ef 1047297cient maizefor Africa (WEMA) Afr Agric Technol Foundation
Courtois BShen L Petalcorin WCarandang S MauleonR LiZ Locating QTLs controllingconstitutive root traits in the rice population IAC 165-Co39 Euphytica 2003134335ndash45
Da-hong L Hui L Yang YL Ping-ping Z Jian-sheng L Down-regulated expression of RACK1geneby RNAinterference enhancesdroughttolerancein rice RiceSci 200916(1)14ndash20
Dalal M Tayal D Chinnusamy V Bansala KC Abiotic stress and ABA-inducible group 4 LEAfrom Brassicanapus plays a key rolein saltand droughttolerance J Biotechnol 2009139137ndash45
Ding Z Li S An X Liu X Qin H Wang D Transgenic expression of MYB15 confers enhancedsensitivity to abscisic acid and improved drought tolerance in Arabidopsis thaliana
J Genet Genomics 20093617ndash29Dubouzet JG Sakuma Y Ito Y Kasuga M Dubouzet EG Miura S et al OsDREB genes in
rice Oryza sativa L encode transcription activators that function in drought high-salt- and cold-responsive gene expression Plant J 200333751ndash63
El-Far IA Allan AY Responses of some wheat cultivars to sowing methods and droughtat different stages of growth Assuit J Agric Sci 199526(1)267ndash77
Eltayeb AE Kawano N Badawi GH Kaminaka H Sanekata T Shibahara T et alOverexpression of monodehydroascorbate reductase in transgenic tobacco confersenhanced tolerance to ozonesalt and polyethyleneglycol stresses Planta 2007225(5)1255ndash64
Falconer DS Introduction to quantitative genetics London New York Longman 1989Feng-ling FU Zhi-Lei F Shi-bing G Shu-feng Z Wan-chen L Evaluation and quantitative
inheritanceof several drought-relativetraits in maize Agric SciChina 20087(3)280ndash90Finkelstein R Gampala S Rock C Abscisic acid signaling in seeds and seedlings Plant
Cell 20021415ndash45Flowers TJ Improving crop salt tolerance J Exp Bot 200455307ndash19Fujita Y Fujita M Satoh R Maruyama K Parvez MM Seki M et al AREB1 is a transcription
Giraudat J Parcy F Bertauche N Gosti F Leung J Morris PC et al Current advances inabscisic acid action and signalling Plant Mol Biol 1994261557ndash77
Gorbalenya AE Koonin EV Helicases amino acid sequence comparisons and structurendashfunction relationships Curr Opin Struct Biol 19933419ndash29
Gosal SS Wani SH Kang MS Biotechnology and drought tolerance J Crop Improvement20092319ndash54
Gubis J Vaňkovaacute R Červenaacute V Draguacuteňovaacute M Hudcovicovaacute M Lichtnerovaacute H et alTransformedtobacco plants with increased toleranceto droughtSouthAfr J Bot200773505ndash11
GuoO Zhang J GaoQ Xing SLi F Wang WDrought tolerancethrough over-expressionof mono ubiquitin in transgenic tobacco J Plant Physiol 20081651745 ndash55
Haley SD Johnson JJ Peairs FB Quick JS Stromberger JA Clayshulte SR et al Registration of lsquoRipperrsquo wheat J Plant Registr 200711ndash6
Han SE Park SR Kwon HB Yi BY Lee GB Byun MO Genetic engineering of drought-resistant tobacco plants by introducingthe trehalose phosphorylase (TP) gene fromPleurotus sajor-caju Plant Cell Tissue Organ Cult 200582151ndash8
Harris K Klein R Mullet J Sorghum stay-green QTL individually reduces post-1047298oweringdrought-induced leaf senescence J Exp Bot 200758327ndash38
Hong BS Zong-Suo L Ming-An S LEA proteins in higher plants structure functiongene expression and regulation Colloids Surf B Biointerf 200545131ndash5
Howarth CJ Yadav RS Successful marker assisted selection for drought tolerance anddisease resistance in pearl milletIGER Innovations 2002
Humphreys MO Humphreys MW Breeding for stress resistance general principles InAshraf M Harris PJC editors Abiotic stresses plant resistance through breedingand molecular approaches 2005 p 19ndash46
Ingram J Bartels D The molecular basis of dehydration tolerance in plants Ann RevPlant Physiol Plant Mol Biol 199647377ndash403
Jaglo-Ottosen KR Gilmour SJ Zarka DG Schabenberger O Thomashow MF ArabidopsisCBF1 overexpression induces COR genes and enhances freezing tolerance Science1998280104ndash6
Jang IC Oh SJ Seo JS Choi WB Song SI Kim CH et al Expression of a bifunctional fusion of the Escherichia coli genes for trehalose-6-phosphate synthase and trehalose-6-phosphate phosphatase in transgenic rice plants increases trehalose accumulationandabiotic stresstolerancewithoutstuntinggrowth Plant Physiol2003131516ndash24
Jeanneau M Gerentes D Foueillassar X Zivy M Vidal J Toppan A et al Improvement of drought tolerance in maize towards the functional validation of the Zm-Asr1 gene andincrease of water use ef 1047297ciency by over-expressing C4-PEPC Biochimie 2002841127ndash35
Jenson KB Palazzo AJ Waldron BL Bushman BS Registration of lsquoFirstStrikersquo Slenderwheatgrass J Plant Registr 2007124ndash5
Joung-youn K Hyung-in C Min-young I Soo-young K Arabidopsis basic leucine zipperproteins that mediate stress-responsive abscisic acid signaling Plant Cell 20024343ndash57
Juskiw PE Helm JH Oro M Nyachiro JM Salmon DF Registration of lsquoBentleyrsquo barley
J Plant Registr 20093(2)119ndash23Kalamaki MS Alexandrou D Lazari D Merkouropoulos G Fotopoulos V Pateraki I et al
Over-expression of a tomato N-acetyl-L -glutamate synthase gene (SlNAGS1) in Arabidopsis thaliana results in high ornithine levels and increased tolerance in saltand drought stresses J Exp Bot 200960(6)1859ndash71
Kamoshita A Babu CR Boopathi NM Fukai S Phenotypic and genotypic analysis of drought-resistance traits for development of rice cultivars adapted to rainfedenvironments Field Crops Res 2008109(103)1-23 doi101016jfcr200806010
KarakasB Ozias-AkinsP Stushnoff C SuefferheldM Rieger M Salinityand drought toleranceof mannitol-accumulating transgenic tobacco Plant Cell Environ 199720609ndash16
Karim S Aronsson H Ericson H Pirhonen M Leyman B Welin B et al Improved droughttolerance without undesired side effects in transgenic plants producing trehalose PlantMol Biol 200764371ndash86
Kasuga M Liu Q Miura S Yamaguchi-Shinozaki K Shinozaki K Improving plant droughtsaltand freezing tolerance by gene transfer of a single stress-inducible transcription factorNat Biotechnol 199917287ndash91
Kindiger M Gaub H Hasegawac M Katsurab Y Ueyamad K Gotob S et al Registrationof lsquoNanryorsquo tall fescue Crop Sci 2006461815ndash6
Kong I Dong J HartGE Characteristics linkage mappositionsand allelic differentiationof Sorghum bicolour (L) Moench by DNA simple-sequence repeats (SSRs) TheorAppl Genet 2000101438ndash48
Kumar R Venuprasad R Atlin GN Genetic analysis of rainfed lowland rice droughttolerance under naturally-occurring stress in eastern India heritability and QTL effects Field Crops Res 200710342ndash52
La1047297tte HR Price AH Courtois B Yield response to water de1047297cit in an upland ricemapping population associations among traits and genetic markers Theor ApplGenet 20041091237ndash46
Lal S Gulyani V Khurana P Overexpressionof HVA1 gene frombarley generates tolerance tosalinity and water stress in transgenic mulberry (Morus indica) Transgenic Res 200817
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63Lanceras J Pantuwan G Jongdee B Toojinda T Quantitative trait loci associated withdrought tolerance at reproductive stage in rice Plant Physiol 2004135384ndash99
Levi A Ovnat L Paterson AH Saranga Y Photosynthesis of cotton near-isogenic linesintrogressed with QTLs for productivity and drought related traits Plant Sci 2009a17788ndash96
Levi A Paterson AH Barak V Yakir D Wang B Chee PW et al Field evaluation of cottonnear-isogenic lines introgressed with QTLs for productivity and drought relatedtraits Mol Breed 2009b23179ndash95
Liu Q Kasuga M Sakuma Y Abe H Miura S Yamaguchi- Shinozaki K et al Twotranscription factors DREB1 and DREB2 with an EREBPAP2 DNA binding domainseparate two cellular signal transduction pathways in drought-and low-temper-ature-responsive gene expression respectively in Arabidopsis Plant Cell 1998101391ndash406
Liu X Hua X Guo J Qi D Wang L Liu Z et al Enhanced tolerance to drought stress intransgenic tobacco plants overexpressing Biotechnol Lett 2008301275ndash80
Liu X Wanga Z Wanga L Wua R Phillips J Deng X LEA 4 group genes from theresurrection plant Boea hygrometrica confer dehydration tolerance in transgenictobacco Plant Sci 200917690ndash8
Luchi S Kobayashi M Taji T Naramoto M Seki M Kato T et al Regulation of droughttolerance by gene manipulation of 9-cis-epoxycarotenoid dioxygenase a keyenzyme in abscisic acid biosynthesis in Arabidopsis Plant J 200127325ndash33
Luo Y Liu YB Dong YX Gao XQ Zhang YS Expression of a putative alfalfa helicase increasestolerance to abioticstress in Arabidopsis by enhancing the capacities for ROS scavengingand osmotic adjustment J Plant Physiol 2009166385ndash94
MacLean JLDaweDC Hardy B Hettel GPRiceAlmanac sourcebook for themostimportanteconomic activity on Earth3rd ed Wallingford England CABI Publishing 2002
Mansour MMF Nitrogen containing compounds and adaptation of plants to salinitystress Biol Plant 200043491ndash500
McKersie BD Bowley SR Harjanto E Leprince O Water-de1047297cit tolerance and 1047297eldperformance of transgenic alfalfa overexpressing superoxide dismutase Plant Physiol19961111177ndash81
McKersie BD Murnaghan J Bowley SR Manipulating freezing tolerance in transgenicplants Acta Physiol Plant 199719485ndash95
Miranda JA Avonce N Suaacuterez R Thevelein JM Dijck PV Iturriaga G A bifunctional TPS-TPP enzyme from yeast confers tolerance to multiple and extreme abiotic-stressconditions in transgenic Arabidopsis Planta 2007226(6)1411ndash21
Mohammadi M Taleei A Zeinali H Naghavi MR Ceccarelli S Grando Baum M QTL analysis for phenologic traits in doubled haploid population of barley Int J AgricBiol 20057(5)820ndash3
Mohanty A Kathuria H Ferjani A Sakamoto A Mohanty P Murata N et al Transgenicsof an elite indica rice variety Pusa Basmati 1 harbouring the codA gene are highlytolerant to salt stress Theor Appl Genet 200210651ndash7
Noaman MM El Sayad AA Asaad FA El Sherbini AM El Bawab AO El Moselhi MA et alRegistration of lsquoGiza 126rsquo barley Crop Sci 199535(6)1710
Noaman MM Ahmed IA El-Sayed AA Abo-El-Enin RA El-Gamal AS El-Sherbiny AM et alRegistration of lsquoGiza 2000rsquo drought-tolerant six-rowed barley for rainfed and newreclaimed areas in Egypt Crop Sci 200747440
Obert DE Evans CP Wesenberg DM Windes JM Erickson CA Jackson EW et alRegistration of lsquoLenetahrsquo spring barley J Plant Registr 20082(2)85ndash7
Oh SJ Jeong JS Kim EH Yi NR Yi SI Jang IC et al Matrix attachment region fromthe chicken lysozyme locus reduces variability in transgene expression andconfers copy number-dependence in transgenic rice plants Plant Cell Rep20054145ndash54
Oh SJ Kim YS Kwon C Park HK Jeong JS Kim JK Overexpression of the transcriptionfactor AP37 in rice improves grain yield under drought conditions Plant Physiol
20092191ndash
200 doi101104pp109137554Owttrim GW RNA helicases and abiotic stress Nucleic AcidsRes 200634(11)3220ndash30Park BJ Liu Z Kanno A Kameya T Increased tolerance to salt and water de1047297cit stress in
transgenic lettuce (Lactuca sativa L) by constitutive expression of LEA PlantGrowth Regul 2005a45165ndash71
Park BJ Liu Z Kanno A Kameya T Genetic improvement of Chinese cabbage for salt anddroughttolerance by constitutiveexpression of a B napus LEAgenePlantSci 2005b169553ndash8
Pastori GM Foyer CH Common componentsnetworks and pathways of crosstoleranceto stress The central role of ldquoredoxrdquo and abscisic acid-mediated controls PlantPhysiol 2002129460ndash8
PellegrineschiA ReynoldsM PachecoM Brito RMAlmeraya R Yamaguchi-ShinozakiK et alStress-induced expression in wheat of the Arabidopsis thaliana DREB1Agenedelayswaterstress symptoms under greenhouse conditions Genome 200447493ndash500
Perl A Perl-Treves R Galili S Aviv D Shalgi E Malkin S et al Enhanced oxidative-stressdefense in transgenic potato overexpressing tomato Cu Zn superoxide dismutaseTheor Appl Genet 199385568ndash76
Podlich DW Winkler CR Cooper M Mapping as you go an effective approach formarker-assisted selection of complex traits Crop Sci 2004441560ndash71
182 M Ashraf Biotechnology Advances 28 (2010) 169ndash183
Pratt RC Casey MA Registration of maize germplasm line Oh605 Crop Sci 2006461004ndash5Qin F Kakimoto M Sakuma Y Maruyama K Osakabe Y Tran LSP et al Regulation and
functional analysis of ZmDREB2A in response to drought and heat stress in Zea mays LPlant J 20075054ndash69
Quan R Shang M Zhang H Zhao Y Zhang J Engineering of enhanced glycinebetainesynthesis improves drought tolerance in maize Plant Biotechnol J 20042477ndash86
Quarrie SA Gulli M Calestani C Steed A Marmiroli N Location of a gene regulatingdrought-induced abscisic acid production on the long arm of chromosome 5A of wheat Theor Appl Genet 199489794ndash800
QuickJS Stromberger JA Clayshulte S Clifford B Johnson JJ Peairs FB et al Registrationof lsquoPrairie Redrsquo wheat Crop Sci 2001411362ndash3
RajaramS Prospects andpromise ofwheatbreedingin the21stcentury Euphytica 20011193-15Rajaram S Role of conventional plant breeding and biotechnology in future wheat
production Turk J Agric Forest 200529105ndash11Rami JF Dufour P Trouche G Fliedel G Mestres C Davrieux F et al Quantitative trait
loci for grain quality productivity morphological and agronomical traits insorghum (Sorghum bicolor L Moench) Theor Appl Genet 199897605ndash16
Rassaa N Salahb HBH Latiri K Thermal responses of durum wheat Triticum durum toearly water stress consequence on leaf and 1047298ower development Plant Biol Pathol2008331(5)363ndash71
Reddy LJ Nigam SN Rao RCN Reddy NS Registration of ICGV 87354 peanut germplasmwith drought tolerance and rust resistance Crop Sci 200141274ndash5
Ribaut JM Ragot M Marker-assisted selection to improve drought adaptation in maize thebackcross approach perspectives limitations and alternatives J Exp Bot 200658351ndash60
Robin S Pathan MS Courtois B La1047297tte R Carandang S Lanceras S et al Mappingosmotic adjustment in an advanced back-cross inbred population of rice Theor
Appl Genet 2003107(7)1288ndash96Romero C Belles JM Vaya JL Serrano R Culianez-Macia FA Expression of the yeast
trehalose-6-phosphate synthase gene in transgenic tobacco plants pleiotropicphenotypes include drought tolerance Planta 1997201293ndash7
Ronde JAD Cress WA Krugerd GHJ Strasserd RJ Van Staden J Photosynthetic responseof transgenic soybean plants containing an Arabidopsis P5CR gene during heat anddrought stress J Plant Physiol 20041611211ndash24
Salvi S Tuberosa R To clone or not to clone plant QTLs present and future challengesTrends Plant Sci 200510297ndash304
Sanchez AC Subudhi PK Rosenow DT Nguyen HT Mapping QTLs associated with droughtresistance in sorghum (Sorghum bicolor L Moench) Plant Mol Biol 200248713ndash26
Saranga Y Menz M Jiang CX Wright RJ Yakir D Paterson AH Genomic dissection of genotypetimesenvironment interactions conferring adaptation of cotton to aridconditions Genome Res 2001111988ndash95
Sari-Gorla M Krajewski P Di Fonzo N Villa M Frova C Genetic analysis of droughttolerance in maize by molecular markers II Plant height and 1047298owering Theor ApplGenet 199999289ndash95
Seki M Narusaka M Ishida J Nanjo T Fujita M Oono Y et al Monitoring the expressionpro1047297les of 7000 Arabidopsis genes under drought cold and high-salinity stressesusing a full-length cDNA microarray Plant J 200231279ndash92
Serraj R Sinclair TR Osmolyte accumulation can it really increase crop yield underdrought conditions Plant Cell Environ 200225333ndash41
Serraj R Krishnamurthy L Kashiwagi J Kumar J Chandra S Crouch JH Variation in roottraits of chickpea (Cicer arietinum L) grown under terminal drought Field CropsRes 200488115ndash27
Serraj R Hash CT Rizvi MHS Sharma A Yadav RS Bidinger FR Recent advances in marker-assisted selection for drought tolerance in pearl millet Plant Prod Sci 20058(3)334ndash7
Shen YG Du BX Zhang WK Zhang JS Chen SY AhCMO regulated by stresses in Atriplexhortensis can improve drought tolerance in transgenic tobacco Theor Appl Genet2002105815ndash21
Shinozaki K Yamaguchi-Shinozaki K Molecular responses to dehydration and lowtemperature differences and cross-talk between two stress signaling pathwaysCurr Opin Plant Biol 20003217ndash23
Singh KB Omar M Saxena MC Johansen C Registration of FLIP 87-59C a drought-tolerant chickpea germplasm line Crop Sci 199636(2)1ndash2
Singh SP Teran H Gutierrez JA Registration of SEA 5 and SEA 13 drought tolerant drybean germplasm Crop Sci 200141276ndash7
Singh SP Teran H Lema M Schwartz HF Miklas PN Registration of white moldresistant dry bean germplasm line A 195 J Plant Registr 2007162ndash3
Sivamani E Bahieldin A Wraith JM Improved biomass productivity and water useef 1047297ciency under water de1047297cit conditions in transgenic wheat constitutivelyexpressing the barley HVA1 gene Plant Sci 20001551ndash9
Skovmand B Reynolds MP DeLacy IH Searching genetic resources for physiologicaltraits with potential for increasing yield In Reynolds MP Ortiz-Monasterio IMcNab A editors Application of Physiology in Wheat Breeding 2001 p 17ndash28
Steele K Novel upland rice variety bred using marker-assisted selection and client-oriented breeding released in Jharkhand India Bangor University 2009
Steele KA Price AH Shashidar HE Witcombe JR Marker-assistedselection to introgressrice QTLs controlling root traitsinto an Indianupland rice variety Theor Appl Genet2006112208ndash21
Steele KA Virk DS Kumar R Prasad SC Witcombe JR Field evaluation of upland ricelines selected for QTLs controlling root traits Field Crops Res 2007101180 ndash6
Sunkar R Kapoor A Zhu JK Post transcriptional induction of two CuZn superoxidedismutase genes in Arabidopsis is mediated by down regulation of miR398 andimportant for oxidative stress tolerance Plant Cell 2006182051ndash65
Talame V Sanguineti MC Chiapparino E Bahri H Ben Salem M Forster BP et alIdenti1047297cation of Hordeum spontaneum QTL alleles improving 1047297eld performance of barley grown under rainfed conditions Ann Appl Bot 2004144309ndash20
Tanner NKCordinO BanroquesJ DoereM Linder PThe Q Motif a newlyidenti1047297ed motif in DEAD box helicases may regulate ATP binding and hydrolysis Mol Cell 200311127ndash38
Taramino G Tarchini R Ferrario S Lee M Pe ME Characterization and mapping of simplesequence repeats (SSRs) in Sorghum bicolor Theor Appl Genet 19979566ndash72
Teulat B Monneveux P Wery J Borriegraves C Souyris I Charrier A et al Relationshipsbetween relative water content andgrowth parameters in barleya QTLstudy NewPhytol 199713799-107
Thi Lang N Chi Buu B Fine mapping for drought tolerance in rice ( Oryza sativa L)Omonrice 2008169-15Till BJ Comai L Henikoff S Tillering and ecotillering for crop improvement In
Varshney RK Tuberosa R editors Genomics-assisted crop improvement genomicsapproaches and platforms vol 1 2007 p 333ndash50
Tondelli A Francia E Barabaschi D Aprile A Skinner JS Stockinger EJ et al Mappingregulatory genes as candidates for cold and drought stress tolerance in barleyTheor Appl Genet 2006112445ndash54
Tuberosa R Salvi S Genomics approaches to improve drought tolerance in cropsTrends Plant Sci 200611405ndash12
Valkoun JJ Wheat pre-breeding using wild progenitors Euphytica 200111917ndash23Vashisht AA Tuteja N Stress responsive DEAD-box helicases a new pathway to
engineer plant stress tolerance J Photochem Photobiol B Biol 200684150 ndash60Vendruscolo ECG Schuster I Pileggi M Scapim CA Molinari HBC Marur CJ et al Stress-
induced synthesis of proline confers tolerance to water de1047297cit in transgenic wheat J Plant Physiol 20071641367ndash76
Vienne D Leonardi A Damerval C Zivy M Genetics of proteome variation for QTL characterization application to drought stress responses in maize J Exp Bot 199950303ndash9
Villareal RL Mujeeb-Kazi A Rajaram S Toro ED Morphological variability in somesynthetic hexaploid wheats derived from Triticum turgidum times T tauschii J GenetBreed 1994487-16
Vinh NT Paterson AH Genome mapping and its implication for stress resistance inplants In Ashraf M Harris PJC editors Abiotic stresses plant resistance throughbreeding and molecular approaches 2005
Vinocur B Altman A Recent advances in engineering plant tolerance to abiotic stressachievements and limitations Curr Opin Biotechnol 200516123ndash32
Wang YJ Hao YJ Zhang ZG Chen T Zhang JS Chen SY Isolation of trehalose-6-phosphate phosphatase gene from tobacco and its functional analysis in yeast cells
J Plant Physiol 2005162215ndash23Wu R Garg A Engineering rice plants with trehalose-producing genes improves
tolerance to drought salt and low temperature ISB News Report 2003Xiao B Huang Y Tang N Xiong L Overexpression of LEA gene in rice improves drought
resistance under 1047297eld conditions Theor Appl Genet 200711535ndash46Xinglai P Sangang X Qiannying P Yinhong S Registration of lsquo Jinmai 50rsquo wheat Crop Sci
200646983ndash5Xiong L Lee H Ishitani M Zhu JK Regulation of osmotic stress-responsive gene
expression by the LOS6ABA1 locus in Arabidopsis J Biol Chem 20022778588ndash96Xu GW Magill CW Shertz KF Hart GE A RFLP linkage map of Sorghum bicolor (L)
Moench Theor Appl Genet 199489139ndash45Xu D Duan X Wang B Hong BHo THD Wu RExpression of a late embryogenesis abundant
protein gene HVA1 from barley confers tolerance to water de1047297cit and salt stress intransgenic rice Plant Physiol 1996110249ndash57
Yadav RS Hash CT Bidinger FR Devos KM Howarth CJ Genomic regions associatedwith grain yield and aspects of post-1047298owering drought tolerance in pearl milletacross stress environments and testers background Euphytica 2004136265ndash77
YamadaM MorishitaH Urano K Shiozaki N Yamaguchi-Shinozaki K ShinozakiK et alEffects of free proline accumulation in petunias under drought stress J Exp Bot2005561975ndash81
Yang WJ Rich PJ Axtell JD Wood KV Bonham CC Ejeta G et al Genotypic variation forglycinebetaine in sorghum Crop Sci 200343162ndash9
Yan-Ying QU Ping MU Xue-Qin L Yu-Xiu T Feng W Hong-Liang Z et al QTL mappingand correlations between leaf water potential and drought resistance in rice underupland and lowland environments Acta Agron Sin 200834(2)198ndash206
Ye XDWu XLZhaoH Frehner MNoumlsberger J Potrykus Iet al Altered fructan accumulationin transgenic Lolium multi 1047298orum plants expressing a Bacillus subtilis sacB gene Plant Cell
Rep 200120205ndash
12Zaharieva M Gaulin E Havaux M Acevedo E Monneveux P Drought and heat responses in
the wild wheat relative Aegilops geniculata Roth potential interest for wheatimprovement Crop Sci 2001411321ndash9
Zhang J Zheng HGAartiA Pantuwan G NguyenTT Tripathy JNet alLocatinggenomicregions associated with components of drought resistance in rice comparativemapping within and across species Theor Appl Genet 200110319ndash29
Zhang GH Su Q An LJ Wu S Characterization and expression of a vacuolar Na+H+
antiporter gene from the monocot halophyte Aeluropus littoralis Plant PhysiolBiochem 200846117ndash26
Zhao J QTLs for oil content and their relationships to other agronomic traits in anEuropeantimesChinese oilseed rape population Germany Diss Grorg-Agust Univer-sity of Goettingen 2002
Zhao J Ren W Zhi D Wang L Xia G Arabidopsis DREB1ACBF3 bestowed transgenic tallfescue increased tolerance to drought stress Plant Cell Rep 2007261521ndash8
Zhou W Li Y Zhao BC Ge RC Shen YZ Wang G Huang ZJ Over-expression of TaSTRG gene improves salt and drought tolerance in rice J Plant Physiol 200915(166)1660ndash70
and cost-effective Molecular mapping and analysis of QTL have been
carried out for a number of qualitative and quantitative traits including
stress tolerance which has undoubtedly resulted in a great magnitude
of knowledge and better understanding of the causal genetic phenom-
ena that regulate these traits However limited success has resulted by
using this knowledge to manipulate genes in an effective way for the
improvement of a speci1047297c trait in crops although some crop cultivars
developed through using these tools have performed extremely well
under1047297
eld stressconditions However there are a variety of reasons forthe limited success in terms of achieving enhanced drought tolerance
using the marker-assisted breeding approach For example a major
dif 1047297culty confronting the scientists is the challenge of precise QTL
identi1047297cation In addition a substantial genetictimes environment interac-
tion inconsistent repeatability large number of genes regulating yield
and invalid use of mapping populations have hampered the pursuits
involving mapping of QTL for enhanced drought stress tolerance
Despite these other factors also impede the application of QTL for
genetic improvement of a trait For example due to unfavorable
epistatic interaction it is hard to transfer the effects of a desired allele to
an elite background material (Podlich et al 2004 Collins et al 2008)
Furthermore in some cases QTL from a speci1047297c background do not
show signi1047297cant effects or cease completely in different backgrounds
even under analogous growth conditions (Cho and Hong 2006 Collins
et al 2008) This makes the utilization of QTL more intricate
Transformation of the knowledge acquired from QTL-oriented molec-
ular studies into a well-de1047297ned upshot for the stakeholders is one of the
key challenges confronting the breeders Certainly a multidisciplinary
approach including more speci1047297cally the identi1047297cation of QTL mediated
signal transduction in response to stresses needs to be adopted
Furthermore although QTL cloning procedures are unwieldy they can
lead to a meaningful outcome because a cloned QTL can offer a reliable
markerfor MABas well as furnish an outlinefor thedetection of superior
allelic variants in crop species (Till et al 2007)
Genetic engineering (Transgenic approach) offers a promise whereby
one can expect a substantial improvement in a desired trait within the
shortest stretch of time Engineering genes encoding organic osmolytes
plant growth regulators antioxidants late embryogenesis abundant pro-
teins and transcription factors has resulted into transgenic lines whichhave out-performed under controlled stress conditions In most cases the
claims made by the researchers in terms of performance of transgenic
lines tested under controlled conditions are considerably in1047298ated
However with the exception of a very few notable cases most of the
transgenic lines of differentcrops have been rarely1047297eld-tested Thushow
far these transgenic lines perform under 1047297eld stress conditions is not
known because under natural1047297eld conditions a linecultivar has to face a
multitude of environmental factors other than the drought stress Thus
appraisal of performance of a transgenic line under natural 1047297eld con-
ditionsis mandatory forthe stakeholdersperspective It is also imperative
to note that most of the drought tolerant transgenic lines have been
developed using a single gene transformation which may not be as
productive as if it had been developed using transformation of many
genes Thus transferring a number of prominent genes effectively in-volved in stress tolerance to transgenic plants seems to be a logical
approach Although a large number of genes appear to be involved in
stress tolerance and most of them have been fully characterized the
function of many of them in the mechanism of stress tolerance is yet to be
investigated
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Ashraf M Breeding for salinity tolerance in plants Crit Rev Plant Sci 19941317 ndash42Ashraf M Akram NA Improving salinity tolerance of plants through conventional
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Ashraf M Foolad MR Roles of glycinebetaine and proline in improving plant abioticstress resistance Environ Exp Bot 200759206ndash16
Ashraf M Athar HR Harris PJC Kwon TR Some prospective strategies for improvingcrop salt tolerance Adv Agron 20089745-110
Asins MJ Present and future of quantitative trait locus analysis in plant breeding PlantBreed 2002121281ndash91
Babu RC Nguyen BD Chamarerk V Shanmugasundaram P Chezhian P Jeyaprakash P et alGeneticanalysis of droughtresistance in riceby molecular markers association betweensecondary traits and 1047297eld performance Crop Sci 2003431457ndash69
Babu RC Zhang J Blum A Ho THD Wu R Nguyen HT HVA1 a LEA gene from barleyconfers dehydration tolerance in transgenic rice( Oryzasativa L)via cell membraneprotection Plant Sci 2004166855ndash62
Badawi GH Kawano N Yamauchi Y Over-expression of ascorbate peroxidase in
tobacco chloroplasts enhances the tolerance to salt stress and water de1047297cit PhysiolPlant 2004121231ndash8Badu-Apraku B Yallou CG Registration of striga-resistant and drought tolerant tropical
early maize populations TZE-W Pop DT STR C4 and TZE-Y Pop DT STR C4 J PlantRegistr 20093(1)86ndash90
Badu-Apraku B Menkir A Kling JG Fakorede MAB Registration of 16 striga resistantearly maturing tropical maize inbred lines Crop Sci 2006461410ndash1
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Baumlnziger M Setimela PS Hodson D Vivek B Breeding for improved drought tolerance inmaize adapted to southern Africa Proceedings of the 4th International Crop ScienceCongress Brisbane Australia Published on CDROM 2004 26 Sep ndash 1 Oct
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traits in the Mediterranean environment identi1047297ed in recombinant inbred lines of the cross lsquoArtarsquo H spontaneum 41-1 Theor Appl Genet 20031071215ndash25
Bergman JW Riveland NR Flynn CR Carlson GR Wichman DM Registration of lsquoMorlinrsquo
saf 1047298ower Crop Sci 2001411640Bernier J Kumar A Venuprasad R Spaner D Atlin G A large-effect QTL for
grain yield under reproductive-stage drought stress in upland rice Crop Sci200747507ndash18
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Bernier J Serraj R Kumar A Venuprasad R Impa S Gowdaa RPV et al The large-effectdrought-resistance QTL qtl121 increases water uptake in upland rice Field CropsRes 200911039ndash46
Bhatnagar-Mathur P ReddyDS Lavanya M Yamaguchi-Shinozaki K Sharma KK Stress-inducible expression of Arabidopsis thaliana DREB1A in transgenic peanut ( Arachishypogaea L) increases transpiration ef 1047297ciency under water-limiting conditionsPlant Cell Rep 2007262071ndash82
Bhatnagar-Mathur P Devi MJ Vadez V Sharma KK Differential antioxidative responses intransgenic peanut bear no relationship to their superior transpiration ef 1047297ciency underdrought stress J Plant Physiol 2009166(11)1207ndash17 doi101016jjplph200901001
Bidinger FR Serraj R Rizvi SMH Howarth C Yadav RS Hash CT Field evaluation of droughttolerance QTL effects on phenotype and adaptation in pearl millet [ Pennisetum glaucum(L) R Br] topcross hybrids Field Crops Res 200594(1)14ndash32
Bidinger FR Nepolean T Hash CT Yadav RS Howarth CJ Identi1047297cation of QTLs for grainyield of pearl millet (Pennisetum glaucum (L) R Br) in environments with variablemoisture during grain 1047297lling Crop Sci 200747969ndash80
Bowers JE Abbey C Anderson S Chang C Draye X A high-density geneticrecombination map of sequence-tagged sites for Sorghum as a framework forcomparative structural and evolutionary genomics of tropical grains and grassesGenetics 2003165367ndash86
Brick MA Ogg JB Singh SP Schwartz HF Johnson JJ Pastor-Corrales MA Registration of drought-tolerant rust-resistant high-yielding pinto bean germplasm lineCO46348 J Plant Registr 20082(2)120ndash4
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Busk PK Pages M Regulation of abscisic acid induced transcription Plant Mol Biol199837425ndash35
Carena MJ Wanner DW Development of genetically broad-based inbred lines of maizefor early-maturing (70-80RM) hybrids J Plant Registr 20093107ndash11
Cash SDBrucknerPL Wichman DMKephart KD Berg JEBoynerR et alRegistration of Willow Creek forage wheat J Plant Registr 20093(2)185ndash90
Cattivelli L Rizza F Badeck FW Mazzucotelli E Francia AMEM Mare AT et al Droughttolerance improvement in crop plants an integrated view from breeding to genomics
Field Crops Res 20081051-14Chen M Wang QY Cheng XG Xu ZS Li LC Ye XG et al GmDREB2 a soybean DRE-
binding transcription factor conferred drought and high-salt tolerance intransgenic plants Biochem Biophys Res Commun 2007353299ndash305
Cheng Z Targolli J Huang X Wu R Wheat LEA genes PMA80 and PMA1959 enhancedehydration tolerance of transgenic rice (Oryza sativa L) Mol Breed 20021071ndash82
ChoEK Hong ChB Over-expression of tobacco NtHSP70-1 contributes to drought-stresstolerance in plants Plant Cell Reports 200625349ndash58
Close TJ Dehydrins a commonality in the response of plants to dehydration and lowtemperature Physiol Plant 1997100291ndash6
Collins NC Tardieu F Tuberosa R Quantitative trait loci and crop performance underabiotic stress where do we stand Plant Physiol 2008147469ndash86
Concept Note Combining breeding and biotechnology to develop water ef 1047297cient maizefor Africa (WEMA) Afr Agric Technol Foundation
Courtois BShen L Petalcorin WCarandang S MauleonR LiZ Locating QTLs controllingconstitutive root traits in the rice population IAC 165-Co39 Euphytica 2003134335ndash45
Da-hong L Hui L Yang YL Ping-ping Z Jian-sheng L Down-regulated expression of RACK1geneby RNAinterference enhancesdroughttolerancein rice RiceSci 200916(1)14ndash20
Dalal M Tayal D Chinnusamy V Bansala KC Abiotic stress and ABA-inducible group 4 LEAfrom Brassicanapus plays a key rolein saltand droughttolerance J Biotechnol 2009139137ndash45
Ding Z Li S An X Liu X Qin H Wang D Transgenic expression of MYB15 confers enhancedsensitivity to abscisic acid and improved drought tolerance in Arabidopsis thaliana
J Genet Genomics 20093617ndash29Dubouzet JG Sakuma Y Ito Y Kasuga M Dubouzet EG Miura S et al OsDREB genes in
rice Oryza sativa L encode transcription activators that function in drought high-salt- and cold-responsive gene expression Plant J 200333751ndash63
El-Far IA Allan AY Responses of some wheat cultivars to sowing methods and droughtat different stages of growth Assuit J Agric Sci 199526(1)267ndash77
Eltayeb AE Kawano N Badawi GH Kaminaka H Sanekata T Shibahara T et alOverexpression of monodehydroascorbate reductase in transgenic tobacco confersenhanced tolerance to ozonesalt and polyethyleneglycol stresses Planta 2007225(5)1255ndash64
Falconer DS Introduction to quantitative genetics London New York Longman 1989Feng-ling FU Zhi-Lei F Shi-bing G Shu-feng Z Wan-chen L Evaluation and quantitative
inheritanceof several drought-relativetraits in maize Agric SciChina 20087(3)280ndash90Finkelstein R Gampala S Rock C Abscisic acid signaling in seeds and seedlings Plant
Cell 20021415ndash45Flowers TJ Improving crop salt tolerance J Exp Bot 200455307ndash19Fujita Y Fujita M Satoh R Maruyama K Parvez MM Seki M et al AREB1 is a transcription
Giraudat J Parcy F Bertauche N Gosti F Leung J Morris PC et al Current advances inabscisic acid action and signalling Plant Mol Biol 1994261557ndash77
Gorbalenya AE Koonin EV Helicases amino acid sequence comparisons and structurendashfunction relationships Curr Opin Struct Biol 19933419ndash29
Gosal SS Wani SH Kang MS Biotechnology and drought tolerance J Crop Improvement20092319ndash54
Gubis J Vaňkovaacute R Červenaacute V Draguacuteňovaacute M Hudcovicovaacute M Lichtnerovaacute H et alTransformedtobacco plants with increased toleranceto droughtSouthAfr J Bot200773505ndash11
GuoO Zhang J GaoQ Xing SLi F Wang WDrought tolerancethrough over-expressionof mono ubiquitin in transgenic tobacco J Plant Physiol 20081651745 ndash55
Haley SD Johnson JJ Peairs FB Quick JS Stromberger JA Clayshulte SR et al Registration of lsquoRipperrsquo wheat J Plant Registr 200711ndash6
Han SE Park SR Kwon HB Yi BY Lee GB Byun MO Genetic engineering of drought-resistant tobacco plants by introducingthe trehalose phosphorylase (TP) gene fromPleurotus sajor-caju Plant Cell Tissue Organ Cult 200582151ndash8
Harris K Klein R Mullet J Sorghum stay-green QTL individually reduces post-1047298oweringdrought-induced leaf senescence J Exp Bot 200758327ndash38
Hong BS Zong-Suo L Ming-An S LEA proteins in higher plants structure functiongene expression and regulation Colloids Surf B Biointerf 200545131ndash5
Howarth CJ Yadav RS Successful marker assisted selection for drought tolerance anddisease resistance in pearl milletIGER Innovations 2002
Humphreys MO Humphreys MW Breeding for stress resistance general principles InAshraf M Harris PJC editors Abiotic stresses plant resistance through breedingand molecular approaches 2005 p 19ndash46
Ingram J Bartels D The molecular basis of dehydration tolerance in plants Ann RevPlant Physiol Plant Mol Biol 199647377ndash403
Jaglo-Ottosen KR Gilmour SJ Zarka DG Schabenberger O Thomashow MF ArabidopsisCBF1 overexpression induces COR genes and enhances freezing tolerance Science1998280104ndash6
Jang IC Oh SJ Seo JS Choi WB Song SI Kim CH et al Expression of a bifunctional fusion of the Escherichia coli genes for trehalose-6-phosphate synthase and trehalose-6-phosphate phosphatase in transgenic rice plants increases trehalose accumulationandabiotic stresstolerancewithoutstuntinggrowth Plant Physiol2003131516ndash24
Jeanneau M Gerentes D Foueillassar X Zivy M Vidal J Toppan A et al Improvement of drought tolerance in maize towards the functional validation of the Zm-Asr1 gene andincrease of water use ef 1047297ciency by over-expressing C4-PEPC Biochimie 2002841127ndash35
Jenson KB Palazzo AJ Waldron BL Bushman BS Registration of lsquoFirstStrikersquo Slenderwheatgrass J Plant Registr 2007124ndash5
Joung-youn K Hyung-in C Min-young I Soo-young K Arabidopsis basic leucine zipperproteins that mediate stress-responsive abscisic acid signaling Plant Cell 20024343ndash57
Juskiw PE Helm JH Oro M Nyachiro JM Salmon DF Registration of lsquoBentleyrsquo barley
J Plant Registr 20093(2)119ndash23Kalamaki MS Alexandrou D Lazari D Merkouropoulos G Fotopoulos V Pateraki I et al
Over-expression of a tomato N-acetyl-L -glutamate synthase gene (SlNAGS1) in Arabidopsis thaliana results in high ornithine levels and increased tolerance in saltand drought stresses J Exp Bot 200960(6)1859ndash71
Kamoshita A Babu CR Boopathi NM Fukai S Phenotypic and genotypic analysis of drought-resistance traits for development of rice cultivars adapted to rainfedenvironments Field Crops Res 2008109(103)1-23 doi101016jfcr200806010
KarakasB Ozias-AkinsP Stushnoff C SuefferheldM Rieger M Salinityand drought toleranceof mannitol-accumulating transgenic tobacco Plant Cell Environ 199720609ndash16
Karim S Aronsson H Ericson H Pirhonen M Leyman B Welin B et al Improved droughttolerance without undesired side effects in transgenic plants producing trehalose PlantMol Biol 200764371ndash86
Kasuga M Liu Q Miura S Yamaguchi-Shinozaki K Shinozaki K Improving plant droughtsaltand freezing tolerance by gene transfer of a single stress-inducible transcription factorNat Biotechnol 199917287ndash91
Kindiger M Gaub H Hasegawac M Katsurab Y Ueyamad K Gotob S et al Registrationof lsquoNanryorsquo tall fescue Crop Sci 2006461815ndash6
Kong I Dong J HartGE Characteristics linkage mappositionsand allelic differentiationof Sorghum bicolour (L) Moench by DNA simple-sequence repeats (SSRs) TheorAppl Genet 2000101438ndash48
Kumar R Venuprasad R Atlin GN Genetic analysis of rainfed lowland rice droughttolerance under naturally-occurring stress in eastern India heritability and QTL effects Field Crops Res 200710342ndash52
La1047297tte HR Price AH Courtois B Yield response to water de1047297cit in an upland ricemapping population associations among traits and genetic markers Theor ApplGenet 20041091237ndash46
Lal S Gulyani V Khurana P Overexpressionof HVA1 gene frombarley generates tolerance tosalinity and water stress in transgenic mulberry (Morus indica) Transgenic Res 200817
(4)651ndash
63Lanceras J Pantuwan G Jongdee B Toojinda T Quantitative trait loci associated withdrought tolerance at reproductive stage in rice Plant Physiol 2004135384ndash99
Levi A Ovnat L Paterson AH Saranga Y Photosynthesis of cotton near-isogenic linesintrogressed with QTLs for productivity and drought related traits Plant Sci 2009a17788ndash96
Levi A Paterson AH Barak V Yakir D Wang B Chee PW et al Field evaluation of cottonnear-isogenic lines introgressed with QTLs for productivity and drought relatedtraits Mol Breed 2009b23179ndash95
Liu Q Kasuga M Sakuma Y Abe H Miura S Yamaguchi- Shinozaki K et al Twotranscription factors DREB1 and DREB2 with an EREBPAP2 DNA binding domainseparate two cellular signal transduction pathways in drought-and low-temper-ature-responsive gene expression respectively in Arabidopsis Plant Cell 1998101391ndash406
Liu X Hua X Guo J Qi D Wang L Liu Z et al Enhanced tolerance to drought stress intransgenic tobacco plants overexpressing Biotechnol Lett 2008301275ndash80
Liu X Wanga Z Wanga L Wua R Phillips J Deng X LEA 4 group genes from theresurrection plant Boea hygrometrica confer dehydration tolerance in transgenictobacco Plant Sci 200917690ndash8
Luchi S Kobayashi M Taji T Naramoto M Seki M Kato T et al Regulation of droughttolerance by gene manipulation of 9-cis-epoxycarotenoid dioxygenase a keyenzyme in abscisic acid biosynthesis in Arabidopsis Plant J 200127325ndash33
Luo Y Liu YB Dong YX Gao XQ Zhang YS Expression of a putative alfalfa helicase increasestolerance to abioticstress in Arabidopsis by enhancing the capacities for ROS scavengingand osmotic adjustment J Plant Physiol 2009166385ndash94
MacLean JLDaweDC Hardy B Hettel GPRiceAlmanac sourcebook for themostimportanteconomic activity on Earth3rd ed Wallingford England CABI Publishing 2002
Mansour MMF Nitrogen containing compounds and adaptation of plants to salinitystress Biol Plant 200043491ndash500
McKersie BD Bowley SR Harjanto E Leprince O Water-de1047297cit tolerance and 1047297eldperformance of transgenic alfalfa overexpressing superoxide dismutase Plant Physiol19961111177ndash81
McKersie BD Murnaghan J Bowley SR Manipulating freezing tolerance in transgenicplants Acta Physiol Plant 199719485ndash95
Miranda JA Avonce N Suaacuterez R Thevelein JM Dijck PV Iturriaga G A bifunctional TPS-TPP enzyme from yeast confers tolerance to multiple and extreme abiotic-stressconditions in transgenic Arabidopsis Planta 2007226(6)1411ndash21
Mohammadi M Taleei A Zeinali H Naghavi MR Ceccarelli S Grando Baum M QTL analysis for phenologic traits in doubled haploid population of barley Int J AgricBiol 20057(5)820ndash3
Mohanty A Kathuria H Ferjani A Sakamoto A Mohanty P Murata N et al Transgenicsof an elite indica rice variety Pusa Basmati 1 harbouring the codA gene are highlytolerant to salt stress Theor Appl Genet 200210651ndash7
Noaman MM El Sayad AA Asaad FA El Sherbini AM El Bawab AO El Moselhi MA et alRegistration of lsquoGiza 126rsquo barley Crop Sci 199535(6)1710
Noaman MM Ahmed IA El-Sayed AA Abo-El-Enin RA El-Gamal AS El-Sherbiny AM et alRegistration of lsquoGiza 2000rsquo drought-tolerant six-rowed barley for rainfed and newreclaimed areas in Egypt Crop Sci 200747440
Obert DE Evans CP Wesenberg DM Windes JM Erickson CA Jackson EW et alRegistration of lsquoLenetahrsquo spring barley J Plant Registr 20082(2)85ndash7
Oh SJ Jeong JS Kim EH Yi NR Yi SI Jang IC et al Matrix attachment region fromthe chicken lysozyme locus reduces variability in transgene expression andconfers copy number-dependence in transgenic rice plants Plant Cell Rep20054145ndash54
Oh SJ Kim YS Kwon C Park HK Jeong JS Kim JK Overexpression of the transcriptionfactor AP37 in rice improves grain yield under drought conditions Plant Physiol
20092191ndash
200 doi101104pp109137554Owttrim GW RNA helicases and abiotic stress Nucleic AcidsRes 200634(11)3220ndash30Park BJ Liu Z Kanno A Kameya T Increased tolerance to salt and water de1047297cit stress in
transgenic lettuce (Lactuca sativa L) by constitutive expression of LEA PlantGrowth Regul 2005a45165ndash71
Park BJ Liu Z Kanno A Kameya T Genetic improvement of Chinese cabbage for salt anddroughttolerance by constitutiveexpression of a B napus LEAgenePlantSci 2005b169553ndash8
Pastori GM Foyer CH Common componentsnetworks and pathways of crosstoleranceto stress The central role of ldquoredoxrdquo and abscisic acid-mediated controls PlantPhysiol 2002129460ndash8
PellegrineschiA ReynoldsM PachecoM Brito RMAlmeraya R Yamaguchi-ShinozakiK et alStress-induced expression in wheat of the Arabidopsis thaliana DREB1Agenedelayswaterstress symptoms under greenhouse conditions Genome 200447493ndash500
Perl A Perl-Treves R Galili S Aviv D Shalgi E Malkin S et al Enhanced oxidative-stressdefense in transgenic potato overexpressing tomato Cu Zn superoxide dismutaseTheor Appl Genet 199385568ndash76
Podlich DW Winkler CR Cooper M Mapping as you go an effective approach formarker-assisted selection of complex traits Crop Sci 2004441560ndash71
182 M Ashraf Biotechnology Advances 28 (2010) 169ndash183
Pratt RC Casey MA Registration of maize germplasm line Oh605 Crop Sci 2006461004ndash5Qin F Kakimoto M Sakuma Y Maruyama K Osakabe Y Tran LSP et al Regulation and
functional analysis of ZmDREB2A in response to drought and heat stress in Zea mays LPlant J 20075054ndash69
Quan R Shang M Zhang H Zhao Y Zhang J Engineering of enhanced glycinebetainesynthesis improves drought tolerance in maize Plant Biotechnol J 20042477ndash86
Quarrie SA Gulli M Calestani C Steed A Marmiroli N Location of a gene regulatingdrought-induced abscisic acid production on the long arm of chromosome 5A of wheat Theor Appl Genet 199489794ndash800
QuickJS Stromberger JA Clayshulte S Clifford B Johnson JJ Peairs FB et al Registrationof lsquoPrairie Redrsquo wheat Crop Sci 2001411362ndash3
RajaramS Prospects andpromise ofwheatbreedingin the21stcentury Euphytica 20011193-15Rajaram S Role of conventional plant breeding and biotechnology in future wheat
production Turk J Agric Forest 200529105ndash11Rami JF Dufour P Trouche G Fliedel G Mestres C Davrieux F et al Quantitative trait
loci for grain quality productivity morphological and agronomical traits insorghum (Sorghum bicolor L Moench) Theor Appl Genet 199897605ndash16
Rassaa N Salahb HBH Latiri K Thermal responses of durum wheat Triticum durum toearly water stress consequence on leaf and 1047298ower development Plant Biol Pathol2008331(5)363ndash71
Reddy LJ Nigam SN Rao RCN Reddy NS Registration of ICGV 87354 peanut germplasmwith drought tolerance and rust resistance Crop Sci 200141274ndash5
Ribaut JM Ragot M Marker-assisted selection to improve drought adaptation in maize thebackcross approach perspectives limitations and alternatives J Exp Bot 200658351ndash60
Robin S Pathan MS Courtois B La1047297tte R Carandang S Lanceras S et al Mappingosmotic adjustment in an advanced back-cross inbred population of rice Theor
Appl Genet 2003107(7)1288ndash96Romero C Belles JM Vaya JL Serrano R Culianez-Macia FA Expression of the yeast
trehalose-6-phosphate synthase gene in transgenic tobacco plants pleiotropicphenotypes include drought tolerance Planta 1997201293ndash7
Ronde JAD Cress WA Krugerd GHJ Strasserd RJ Van Staden J Photosynthetic responseof transgenic soybean plants containing an Arabidopsis P5CR gene during heat anddrought stress J Plant Physiol 20041611211ndash24
Salvi S Tuberosa R To clone or not to clone plant QTLs present and future challengesTrends Plant Sci 200510297ndash304
Sanchez AC Subudhi PK Rosenow DT Nguyen HT Mapping QTLs associated with droughtresistance in sorghum (Sorghum bicolor L Moench) Plant Mol Biol 200248713ndash26
Saranga Y Menz M Jiang CX Wright RJ Yakir D Paterson AH Genomic dissection of genotypetimesenvironment interactions conferring adaptation of cotton to aridconditions Genome Res 2001111988ndash95
Sari-Gorla M Krajewski P Di Fonzo N Villa M Frova C Genetic analysis of droughttolerance in maize by molecular markers II Plant height and 1047298owering Theor ApplGenet 199999289ndash95
Seki M Narusaka M Ishida J Nanjo T Fujita M Oono Y et al Monitoring the expressionpro1047297les of 7000 Arabidopsis genes under drought cold and high-salinity stressesusing a full-length cDNA microarray Plant J 200231279ndash92
Serraj R Sinclair TR Osmolyte accumulation can it really increase crop yield underdrought conditions Plant Cell Environ 200225333ndash41
Serraj R Krishnamurthy L Kashiwagi J Kumar J Chandra S Crouch JH Variation in roottraits of chickpea (Cicer arietinum L) grown under terminal drought Field CropsRes 200488115ndash27
Serraj R Hash CT Rizvi MHS Sharma A Yadav RS Bidinger FR Recent advances in marker-assisted selection for drought tolerance in pearl millet Plant Prod Sci 20058(3)334ndash7
Shen YG Du BX Zhang WK Zhang JS Chen SY AhCMO regulated by stresses in Atriplexhortensis can improve drought tolerance in transgenic tobacco Theor Appl Genet2002105815ndash21
Shinozaki K Yamaguchi-Shinozaki K Molecular responses to dehydration and lowtemperature differences and cross-talk between two stress signaling pathwaysCurr Opin Plant Biol 20003217ndash23
Singh KB Omar M Saxena MC Johansen C Registration of FLIP 87-59C a drought-tolerant chickpea germplasm line Crop Sci 199636(2)1ndash2
Singh SP Teran H Gutierrez JA Registration of SEA 5 and SEA 13 drought tolerant drybean germplasm Crop Sci 200141276ndash7
Singh SP Teran H Lema M Schwartz HF Miklas PN Registration of white moldresistant dry bean germplasm line A 195 J Plant Registr 2007162ndash3
Sivamani E Bahieldin A Wraith JM Improved biomass productivity and water useef 1047297ciency under water de1047297cit conditions in transgenic wheat constitutivelyexpressing the barley HVA1 gene Plant Sci 20001551ndash9
Skovmand B Reynolds MP DeLacy IH Searching genetic resources for physiologicaltraits with potential for increasing yield In Reynolds MP Ortiz-Monasterio IMcNab A editors Application of Physiology in Wheat Breeding 2001 p 17ndash28
Steele K Novel upland rice variety bred using marker-assisted selection and client-oriented breeding released in Jharkhand India Bangor University 2009
Steele KA Price AH Shashidar HE Witcombe JR Marker-assistedselection to introgressrice QTLs controlling root traitsinto an Indianupland rice variety Theor Appl Genet2006112208ndash21
Steele KA Virk DS Kumar R Prasad SC Witcombe JR Field evaluation of upland ricelines selected for QTLs controlling root traits Field Crops Res 2007101180 ndash6
Sunkar R Kapoor A Zhu JK Post transcriptional induction of two CuZn superoxidedismutase genes in Arabidopsis is mediated by down regulation of miR398 andimportant for oxidative stress tolerance Plant Cell 2006182051ndash65
Talame V Sanguineti MC Chiapparino E Bahri H Ben Salem M Forster BP et alIdenti1047297cation of Hordeum spontaneum QTL alleles improving 1047297eld performance of barley grown under rainfed conditions Ann Appl Bot 2004144309ndash20
Tanner NKCordinO BanroquesJ DoereM Linder PThe Q Motif a newlyidenti1047297ed motif in DEAD box helicases may regulate ATP binding and hydrolysis Mol Cell 200311127ndash38
Taramino G Tarchini R Ferrario S Lee M Pe ME Characterization and mapping of simplesequence repeats (SSRs) in Sorghum bicolor Theor Appl Genet 19979566ndash72
Teulat B Monneveux P Wery J Borriegraves C Souyris I Charrier A et al Relationshipsbetween relative water content andgrowth parameters in barleya QTLstudy NewPhytol 199713799-107
Thi Lang N Chi Buu B Fine mapping for drought tolerance in rice ( Oryza sativa L)Omonrice 2008169-15Till BJ Comai L Henikoff S Tillering and ecotillering for crop improvement In
Varshney RK Tuberosa R editors Genomics-assisted crop improvement genomicsapproaches and platforms vol 1 2007 p 333ndash50
Tondelli A Francia E Barabaschi D Aprile A Skinner JS Stockinger EJ et al Mappingregulatory genes as candidates for cold and drought stress tolerance in barleyTheor Appl Genet 2006112445ndash54
Tuberosa R Salvi S Genomics approaches to improve drought tolerance in cropsTrends Plant Sci 200611405ndash12
Valkoun JJ Wheat pre-breeding using wild progenitors Euphytica 200111917ndash23Vashisht AA Tuteja N Stress responsive DEAD-box helicases a new pathway to
engineer plant stress tolerance J Photochem Photobiol B Biol 200684150 ndash60Vendruscolo ECG Schuster I Pileggi M Scapim CA Molinari HBC Marur CJ et al Stress-
induced synthesis of proline confers tolerance to water de1047297cit in transgenic wheat J Plant Physiol 20071641367ndash76
Vienne D Leonardi A Damerval C Zivy M Genetics of proteome variation for QTL characterization application to drought stress responses in maize J Exp Bot 199950303ndash9
Villareal RL Mujeeb-Kazi A Rajaram S Toro ED Morphological variability in somesynthetic hexaploid wheats derived from Triticum turgidum times T tauschii J GenetBreed 1994487-16
Vinh NT Paterson AH Genome mapping and its implication for stress resistance inplants In Ashraf M Harris PJC editors Abiotic stresses plant resistance throughbreeding and molecular approaches 2005
Vinocur B Altman A Recent advances in engineering plant tolerance to abiotic stressachievements and limitations Curr Opin Biotechnol 200516123ndash32
Wang YJ Hao YJ Zhang ZG Chen T Zhang JS Chen SY Isolation of trehalose-6-phosphate phosphatase gene from tobacco and its functional analysis in yeast cells
J Plant Physiol 2005162215ndash23Wu R Garg A Engineering rice plants with trehalose-producing genes improves
tolerance to drought salt and low temperature ISB News Report 2003Xiao B Huang Y Tang N Xiong L Overexpression of LEA gene in rice improves drought
resistance under 1047297eld conditions Theor Appl Genet 200711535ndash46Xinglai P Sangang X Qiannying P Yinhong S Registration of lsquo Jinmai 50rsquo wheat Crop Sci
200646983ndash5Xiong L Lee H Ishitani M Zhu JK Regulation of osmotic stress-responsive gene
expression by the LOS6ABA1 locus in Arabidopsis J Biol Chem 20022778588ndash96Xu GW Magill CW Shertz KF Hart GE A RFLP linkage map of Sorghum bicolor (L)
Moench Theor Appl Genet 199489139ndash45Xu D Duan X Wang B Hong BHo THD Wu RExpression of a late embryogenesis abundant
protein gene HVA1 from barley confers tolerance to water de1047297cit and salt stress intransgenic rice Plant Physiol 1996110249ndash57
Yadav RS Hash CT Bidinger FR Devos KM Howarth CJ Genomic regions associatedwith grain yield and aspects of post-1047298owering drought tolerance in pearl milletacross stress environments and testers background Euphytica 2004136265ndash77
YamadaM MorishitaH Urano K Shiozaki N Yamaguchi-Shinozaki K ShinozakiK et alEffects of free proline accumulation in petunias under drought stress J Exp Bot2005561975ndash81
Yang WJ Rich PJ Axtell JD Wood KV Bonham CC Ejeta G et al Genotypic variation forglycinebetaine in sorghum Crop Sci 200343162ndash9
Yan-Ying QU Ping MU Xue-Qin L Yu-Xiu T Feng W Hong-Liang Z et al QTL mappingand correlations between leaf water potential and drought resistance in rice underupland and lowland environments Acta Agron Sin 200834(2)198ndash206
Ye XDWu XLZhaoH Frehner MNoumlsberger J Potrykus Iet al Altered fructan accumulationin transgenic Lolium multi 1047298orum plants expressing a Bacillus subtilis sacB gene Plant Cell
Rep 200120205ndash
12Zaharieva M Gaulin E Havaux M Acevedo E Monneveux P Drought and heat responses in
the wild wheat relative Aegilops geniculata Roth potential interest for wheatimprovement Crop Sci 2001411321ndash9
Zhang J Zheng HGAartiA Pantuwan G NguyenTT Tripathy JNet alLocatinggenomicregions associated with components of drought resistance in rice comparativemapping within and across species Theor Appl Genet 200110319ndash29
Zhang GH Su Q An LJ Wu S Characterization and expression of a vacuolar Na+H+
antiporter gene from the monocot halophyte Aeluropus littoralis Plant PhysiolBiochem 200846117ndash26
Zhao J QTLs for oil content and their relationships to other agronomic traits in anEuropeantimesChinese oilseed rape population Germany Diss Grorg-Agust Univer-sity of Goettingen 2002
Zhao J Ren W Zhi D Wang L Xia G Arabidopsis DREB1ACBF3 bestowed transgenic tallfescue increased tolerance to drought stress Plant Cell Rep 2007261521ndash8
Zhou W Li Y Zhao BC Ge RC Shen YZ Wang G Huang ZJ Over-expression of TaSTRG gene improves salt and drought tolerance in rice J Plant Physiol 200915(166)1660ndash70
and cost-effective Molecular mapping and analysis of QTL have been
carried out for a number of qualitative and quantitative traits including
stress tolerance which has undoubtedly resulted in a great magnitude
of knowledge and better understanding of the causal genetic phenom-
ena that regulate these traits However limited success has resulted by
using this knowledge to manipulate genes in an effective way for the
improvement of a speci1047297c trait in crops although some crop cultivars
developed through using these tools have performed extremely well
under1047297
eld stressconditions However there are a variety of reasons forthe limited success in terms of achieving enhanced drought tolerance
using the marker-assisted breeding approach For example a major
dif 1047297culty confronting the scientists is the challenge of precise QTL
identi1047297cation In addition a substantial genetictimes environment interac-
tion inconsistent repeatability large number of genes regulating yield
and invalid use of mapping populations have hampered the pursuits
involving mapping of QTL for enhanced drought stress tolerance
Despite these other factors also impede the application of QTL for
genetic improvement of a trait For example due to unfavorable
epistatic interaction it is hard to transfer the effects of a desired allele to
an elite background material (Podlich et al 2004 Collins et al 2008)
Furthermore in some cases QTL from a speci1047297c background do not
show signi1047297cant effects or cease completely in different backgrounds
even under analogous growth conditions (Cho and Hong 2006 Collins
et al 2008) This makes the utilization of QTL more intricate
Transformation of the knowledge acquired from QTL-oriented molec-
ular studies into a well-de1047297ned upshot for the stakeholders is one of the
key challenges confronting the breeders Certainly a multidisciplinary
approach including more speci1047297cally the identi1047297cation of QTL mediated
signal transduction in response to stresses needs to be adopted
Furthermore although QTL cloning procedures are unwieldy they can
lead to a meaningful outcome because a cloned QTL can offer a reliable
markerfor MABas well as furnish an outlinefor thedetection of superior
allelic variants in crop species (Till et al 2007)
Genetic engineering (Transgenic approach) offers a promise whereby
one can expect a substantial improvement in a desired trait within the
shortest stretch of time Engineering genes encoding organic osmolytes
plant growth regulators antioxidants late embryogenesis abundant pro-
teins and transcription factors has resulted into transgenic lines whichhave out-performed under controlled stress conditions In most cases the
claims made by the researchers in terms of performance of transgenic
lines tested under controlled conditions are considerably in1047298ated
However with the exception of a very few notable cases most of the
transgenic lines of differentcrops have been rarely1047297eld-tested Thushow
far these transgenic lines perform under 1047297eld stress conditions is not
known because under natural1047297eld conditions a linecultivar has to face a
multitude of environmental factors other than the drought stress Thus
appraisal of performance of a transgenic line under natural 1047297eld con-
ditionsis mandatory forthe stakeholdersperspective It is also imperative
to note that most of the drought tolerant transgenic lines have been
developed using a single gene transformation which may not be as
productive as if it had been developed using transformation of many
genes Thus transferring a number of prominent genes effectively in-volved in stress tolerance to transgenic plants seems to be a logical
approach Although a large number of genes appear to be involved in
stress tolerance and most of them have been fully characterized the
function of many of them in the mechanism of stress tolerance is yet to be
investigated
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Ashraf M Breeding for salinity tolerance in plants Crit Rev Plant Sci 19941317 ndash42Ashraf M Akram NA Improving salinity tolerance of plants through conventional
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Ashraf M Foolad MR Roles of glycinebetaine and proline in improving plant abioticstress resistance Environ Exp Bot 200759206ndash16
Ashraf M Athar HR Harris PJC Kwon TR Some prospective strategies for improvingcrop salt tolerance Adv Agron 20089745-110
Asins MJ Present and future of quantitative trait locus analysis in plant breeding PlantBreed 2002121281ndash91
Babu RC Nguyen BD Chamarerk V Shanmugasundaram P Chezhian P Jeyaprakash P et alGeneticanalysis of droughtresistance in riceby molecular markers association betweensecondary traits and 1047297eld performance Crop Sci 2003431457ndash69
Babu RC Zhang J Blum A Ho THD Wu R Nguyen HT HVA1 a LEA gene from barleyconfers dehydration tolerance in transgenic rice( Oryzasativa L)via cell membraneprotection Plant Sci 2004166855ndash62
Badawi GH Kawano N Yamauchi Y Over-expression of ascorbate peroxidase in
tobacco chloroplasts enhances the tolerance to salt stress and water de1047297cit PhysiolPlant 2004121231ndash8Badu-Apraku B Yallou CG Registration of striga-resistant and drought tolerant tropical
early maize populations TZE-W Pop DT STR C4 and TZE-Y Pop DT STR C4 J PlantRegistr 20093(1)86ndash90
Badu-Apraku B Menkir A Kling JG Fakorede MAB Registration of 16 striga resistantearly maturing tropical maize inbred lines Crop Sci 2006461410ndash1
Baenziger PS Beecher B Graybosch RA Ibrahim AMH Baltensperger DD Nelson LA et alRegistration of lsquoNEO1643rsquo wheat J Plant Registr 20082(1)36ndash42
Baumlnziger M Setimela PS Hodson D Vivek B Breeding for improved drought tolerance inmaize adapted to southern Africa Proceedings of the 4th International Crop ScienceCongress Brisbane Australia Published on CDROM 2004 26 Sep ndash 1 Oct
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traits in the Mediterranean environment identi1047297ed in recombinant inbred lines of the cross lsquoArtarsquo H spontaneum 41-1 Theor Appl Genet 20031071215ndash25
Bergman JW Riveland NR Flynn CR Carlson GR Wichman DM Registration of lsquoMorlinrsquo
saf 1047298ower Crop Sci 2001411640Bernier J Kumar A Venuprasad R Spaner D Atlin G A large-effect QTL for
grain yield under reproductive-stage drought stress in upland rice Crop Sci200747507ndash18
BernierJ Kumar A SerrajR SpanerD Atlin G Review breeding uplandrice fordroughtresistance J Sci Food Agric 200888927ndash39
Bernier J Serraj R Kumar A Venuprasad R Impa S Gowdaa RPV et al The large-effectdrought-resistance QTL qtl121 increases water uptake in upland rice Field CropsRes 200911039ndash46
Bhatnagar-Mathur P ReddyDS Lavanya M Yamaguchi-Shinozaki K Sharma KK Stress-inducible expression of Arabidopsis thaliana DREB1A in transgenic peanut ( Arachishypogaea L) increases transpiration ef 1047297ciency under water-limiting conditionsPlant Cell Rep 2007262071ndash82
Bhatnagar-Mathur P Devi MJ Vadez V Sharma KK Differential antioxidative responses intransgenic peanut bear no relationship to their superior transpiration ef 1047297ciency underdrought stress J Plant Physiol 2009166(11)1207ndash17 doi101016jjplph200901001
Bidinger FR Serraj R Rizvi SMH Howarth C Yadav RS Hash CT Field evaluation of droughttolerance QTL effects on phenotype and adaptation in pearl millet [ Pennisetum glaucum(L) R Br] topcross hybrids Field Crops Res 200594(1)14ndash32
Bidinger FR Nepolean T Hash CT Yadav RS Howarth CJ Identi1047297cation of QTLs for grainyield of pearl millet (Pennisetum glaucum (L) R Br) in environments with variablemoisture during grain 1047297lling Crop Sci 200747969ndash80
Bowers JE Abbey C Anderson S Chang C Draye X A high-density geneticrecombination map of sequence-tagged sites for Sorghum as a framework forcomparative structural and evolutionary genomics of tropical grains and grassesGenetics 2003165367ndash86
Brick MA Ogg JB Singh SP Schwartz HF Johnson JJ Pastor-Corrales MA Registration of drought-tolerant rust-resistant high-yielding pinto bean germplasm lineCO46348 J Plant Registr 20082(2)120ndash4
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Busk PK Pages M Regulation of abscisic acid induced transcription Plant Mol Biol199837425ndash35
Carena MJ Wanner DW Development of genetically broad-based inbred lines of maizefor early-maturing (70-80RM) hybrids J Plant Registr 20093107ndash11
Cash SDBrucknerPL Wichman DMKephart KD Berg JEBoynerR et alRegistration of Willow Creek forage wheat J Plant Registr 20093(2)185ndash90
Cattivelli L Rizza F Badeck FW Mazzucotelli E Francia AMEM Mare AT et al Droughttolerance improvement in crop plants an integrated view from breeding to genomics
Field Crops Res 20081051-14Chen M Wang QY Cheng XG Xu ZS Li LC Ye XG et al GmDREB2 a soybean DRE-
binding transcription factor conferred drought and high-salt tolerance intransgenic plants Biochem Biophys Res Commun 2007353299ndash305
Cheng Z Targolli J Huang X Wu R Wheat LEA genes PMA80 and PMA1959 enhancedehydration tolerance of transgenic rice (Oryza sativa L) Mol Breed 20021071ndash82
ChoEK Hong ChB Over-expression of tobacco NtHSP70-1 contributes to drought-stresstolerance in plants Plant Cell Reports 200625349ndash58
Close TJ Dehydrins a commonality in the response of plants to dehydration and lowtemperature Physiol Plant 1997100291ndash6
Collins NC Tardieu F Tuberosa R Quantitative trait loci and crop performance underabiotic stress where do we stand Plant Physiol 2008147469ndash86
Concept Note Combining breeding and biotechnology to develop water ef 1047297cient maizefor Africa (WEMA) Afr Agric Technol Foundation
Courtois BShen L Petalcorin WCarandang S MauleonR LiZ Locating QTLs controllingconstitutive root traits in the rice population IAC 165-Co39 Euphytica 2003134335ndash45
Da-hong L Hui L Yang YL Ping-ping Z Jian-sheng L Down-regulated expression of RACK1geneby RNAinterference enhancesdroughttolerancein rice RiceSci 200916(1)14ndash20
Dalal M Tayal D Chinnusamy V Bansala KC Abiotic stress and ABA-inducible group 4 LEAfrom Brassicanapus plays a key rolein saltand droughttolerance J Biotechnol 2009139137ndash45
Ding Z Li S An X Liu X Qin H Wang D Transgenic expression of MYB15 confers enhancedsensitivity to abscisic acid and improved drought tolerance in Arabidopsis thaliana
J Genet Genomics 20093617ndash29Dubouzet JG Sakuma Y Ito Y Kasuga M Dubouzet EG Miura S et al OsDREB genes in
rice Oryza sativa L encode transcription activators that function in drought high-salt- and cold-responsive gene expression Plant J 200333751ndash63
El-Far IA Allan AY Responses of some wheat cultivars to sowing methods and droughtat different stages of growth Assuit J Agric Sci 199526(1)267ndash77
Eltayeb AE Kawano N Badawi GH Kaminaka H Sanekata T Shibahara T et alOverexpression of monodehydroascorbate reductase in transgenic tobacco confersenhanced tolerance to ozonesalt and polyethyleneglycol stresses Planta 2007225(5)1255ndash64
Falconer DS Introduction to quantitative genetics London New York Longman 1989Feng-ling FU Zhi-Lei F Shi-bing G Shu-feng Z Wan-chen L Evaluation and quantitative
inheritanceof several drought-relativetraits in maize Agric SciChina 20087(3)280ndash90Finkelstein R Gampala S Rock C Abscisic acid signaling in seeds and seedlings Plant
Cell 20021415ndash45Flowers TJ Improving crop salt tolerance J Exp Bot 200455307ndash19Fujita Y Fujita M Satoh R Maruyama K Parvez MM Seki M et al AREB1 is a transcription
Giraudat J Parcy F Bertauche N Gosti F Leung J Morris PC et al Current advances inabscisic acid action and signalling Plant Mol Biol 1994261557ndash77
Gorbalenya AE Koonin EV Helicases amino acid sequence comparisons and structurendashfunction relationships Curr Opin Struct Biol 19933419ndash29
Gosal SS Wani SH Kang MS Biotechnology and drought tolerance J Crop Improvement20092319ndash54
Gubis J Vaňkovaacute R Červenaacute V Draguacuteňovaacute M Hudcovicovaacute M Lichtnerovaacute H et alTransformedtobacco plants with increased toleranceto droughtSouthAfr J Bot200773505ndash11
GuoO Zhang J GaoQ Xing SLi F Wang WDrought tolerancethrough over-expressionof mono ubiquitin in transgenic tobacco J Plant Physiol 20081651745 ndash55
Haley SD Johnson JJ Peairs FB Quick JS Stromberger JA Clayshulte SR et al Registration of lsquoRipperrsquo wheat J Plant Registr 200711ndash6
Han SE Park SR Kwon HB Yi BY Lee GB Byun MO Genetic engineering of drought-resistant tobacco plants by introducingthe trehalose phosphorylase (TP) gene fromPleurotus sajor-caju Plant Cell Tissue Organ Cult 200582151ndash8
Harris K Klein R Mullet J Sorghum stay-green QTL individually reduces post-1047298oweringdrought-induced leaf senescence J Exp Bot 200758327ndash38
Hong BS Zong-Suo L Ming-An S LEA proteins in higher plants structure functiongene expression and regulation Colloids Surf B Biointerf 200545131ndash5
Howarth CJ Yadav RS Successful marker assisted selection for drought tolerance anddisease resistance in pearl milletIGER Innovations 2002
Humphreys MO Humphreys MW Breeding for stress resistance general principles InAshraf M Harris PJC editors Abiotic stresses plant resistance through breedingand molecular approaches 2005 p 19ndash46
Ingram J Bartels D The molecular basis of dehydration tolerance in plants Ann RevPlant Physiol Plant Mol Biol 199647377ndash403
Jaglo-Ottosen KR Gilmour SJ Zarka DG Schabenberger O Thomashow MF ArabidopsisCBF1 overexpression induces COR genes and enhances freezing tolerance Science1998280104ndash6
Jang IC Oh SJ Seo JS Choi WB Song SI Kim CH et al Expression of a bifunctional fusion of the Escherichia coli genes for trehalose-6-phosphate synthase and trehalose-6-phosphate phosphatase in transgenic rice plants increases trehalose accumulationandabiotic stresstolerancewithoutstuntinggrowth Plant Physiol2003131516ndash24
Jeanneau M Gerentes D Foueillassar X Zivy M Vidal J Toppan A et al Improvement of drought tolerance in maize towards the functional validation of the Zm-Asr1 gene andincrease of water use ef 1047297ciency by over-expressing C4-PEPC Biochimie 2002841127ndash35
Jenson KB Palazzo AJ Waldron BL Bushman BS Registration of lsquoFirstStrikersquo Slenderwheatgrass J Plant Registr 2007124ndash5
Joung-youn K Hyung-in C Min-young I Soo-young K Arabidopsis basic leucine zipperproteins that mediate stress-responsive abscisic acid signaling Plant Cell 20024343ndash57
Juskiw PE Helm JH Oro M Nyachiro JM Salmon DF Registration of lsquoBentleyrsquo barley
J Plant Registr 20093(2)119ndash23Kalamaki MS Alexandrou D Lazari D Merkouropoulos G Fotopoulos V Pateraki I et al
Over-expression of a tomato N-acetyl-L -glutamate synthase gene (SlNAGS1) in Arabidopsis thaliana results in high ornithine levels and increased tolerance in saltand drought stresses J Exp Bot 200960(6)1859ndash71
Kamoshita A Babu CR Boopathi NM Fukai S Phenotypic and genotypic analysis of drought-resistance traits for development of rice cultivars adapted to rainfedenvironments Field Crops Res 2008109(103)1-23 doi101016jfcr200806010
KarakasB Ozias-AkinsP Stushnoff C SuefferheldM Rieger M Salinityand drought toleranceof mannitol-accumulating transgenic tobacco Plant Cell Environ 199720609ndash16
Karim S Aronsson H Ericson H Pirhonen M Leyman B Welin B et al Improved droughttolerance without undesired side effects in transgenic plants producing trehalose PlantMol Biol 200764371ndash86
Kasuga M Liu Q Miura S Yamaguchi-Shinozaki K Shinozaki K Improving plant droughtsaltand freezing tolerance by gene transfer of a single stress-inducible transcription factorNat Biotechnol 199917287ndash91
Kindiger M Gaub H Hasegawac M Katsurab Y Ueyamad K Gotob S et al Registrationof lsquoNanryorsquo tall fescue Crop Sci 2006461815ndash6
Kong I Dong J HartGE Characteristics linkage mappositionsand allelic differentiationof Sorghum bicolour (L) Moench by DNA simple-sequence repeats (SSRs) TheorAppl Genet 2000101438ndash48
Kumar R Venuprasad R Atlin GN Genetic analysis of rainfed lowland rice droughttolerance under naturally-occurring stress in eastern India heritability and QTL effects Field Crops Res 200710342ndash52
La1047297tte HR Price AH Courtois B Yield response to water de1047297cit in an upland ricemapping population associations among traits and genetic markers Theor ApplGenet 20041091237ndash46
Lal S Gulyani V Khurana P Overexpressionof HVA1 gene frombarley generates tolerance tosalinity and water stress in transgenic mulberry (Morus indica) Transgenic Res 200817
(4)651ndash
63Lanceras J Pantuwan G Jongdee B Toojinda T Quantitative trait loci associated withdrought tolerance at reproductive stage in rice Plant Physiol 2004135384ndash99
Levi A Ovnat L Paterson AH Saranga Y Photosynthesis of cotton near-isogenic linesintrogressed with QTLs for productivity and drought related traits Plant Sci 2009a17788ndash96
Levi A Paterson AH Barak V Yakir D Wang B Chee PW et al Field evaluation of cottonnear-isogenic lines introgressed with QTLs for productivity and drought relatedtraits Mol Breed 2009b23179ndash95
Liu Q Kasuga M Sakuma Y Abe H Miura S Yamaguchi- Shinozaki K et al Twotranscription factors DREB1 and DREB2 with an EREBPAP2 DNA binding domainseparate two cellular signal transduction pathways in drought-and low-temper-ature-responsive gene expression respectively in Arabidopsis Plant Cell 1998101391ndash406
Liu X Hua X Guo J Qi D Wang L Liu Z et al Enhanced tolerance to drought stress intransgenic tobacco plants overexpressing Biotechnol Lett 2008301275ndash80
Liu X Wanga Z Wanga L Wua R Phillips J Deng X LEA 4 group genes from theresurrection plant Boea hygrometrica confer dehydration tolerance in transgenictobacco Plant Sci 200917690ndash8
Luchi S Kobayashi M Taji T Naramoto M Seki M Kato T et al Regulation of droughttolerance by gene manipulation of 9-cis-epoxycarotenoid dioxygenase a keyenzyme in abscisic acid biosynthesis in Arabidopsis Plant J 200127325ndash33
Luo Y Liu YB Dong YX Gao XQ Zhang YS Expression of a putative alfalfa helicase increasestolerance to abioticstress in Arabidopsis by enhancing the capacities for ROS scavengingand osmotic adjustment J Plant Physiol 2009166385ndash94
MacLean JLDaweDC Hardy B Hettel GPRiceAlmanac sourcebook for themostimportanteconomic activity on Earth3rd ed Wallingford England CABI Publishing 2002
Mansour MMF Nitrogen containing compounds and adaptation of plants to salinitystress Biol Plant 200043491ndash500
McKersie BD Bowley SR Harjanto E Leprince O Water-de1047297cit tolerance and 1047297eldperformance of transgenic alfalfa overexpressing superoxide dismutase Plant Physiol19961111177ndash81
McKersie BD Murnaghan J Bowley SR Manipulating freezing tolerance in transgenicplants Acta Physiol Plant 199719485ndash95
Miranda JA Avonce N Suaacuterez R Thevelein JM Dijck PV Iturriaga G A bifunctional TPS-TPP enzyme from yeast confers tolerance to multiple and extreme abiotic-stressconditions in transgenic Arabidopsis Planta 2007226(6)1411ndash21
Mohammadi M Taleei A Zeinali H Naghavi MR Ceccarelli S Grando Baum M QTL analysis for phenologic traits in doubled haploid population of barley Int J AgricBiol 20057(5)820ndash3
Mohanty A Kathuria H Ferjani A Sakamoto A Mohanty P Murata N et al Transgenicsof an elite indica rice variety Pusa Basmati 1 harbouring the codA gene are highlytolerant to salt stress Theor Appl Genet 200210651ndash7
Noaman MM El Sayad AA Asaad FA El Sherbini AM El Bawab AO El Moselhi MA et alRegistration of lsquoGiza 126rsquo barley Crop Sci 199535(6)1710
Noaman MM Ahmed IA El-Sayed AA Abo-El-Enin RA El-Gamal AS El-Sherbiny AM et alRegistration of lsquoGiza 2000rsquo drought-tolerant six-rowed barley for rainfed and newreclaimed areas in Egypt Crop Sci 200747440
Obert DE Evans CP Wesenberg DM Windes JM Erickson CA Jackson EW et alRegistration of lsquoLenetahrsquo spring barley J Plant Registr 20082(2)85ndash7
Oh SJ Jeong JS Kim EH Yi NR Yi SI Jang IC et al Matrix attachment region fromthe chicken lysozyme locus reduces variability in transgene expression andconfers copy number-dependence in transgenic rice plants Plant Cell Rep20054145ndash54
Oh SJ Kim YS Kwon C Park HK Jeong JS Kim JK Overexpression of the transcriptionfactor AP37 in rice improves grain yield under drought conditions Plant Physiol
20092191ndash
200 doi101104pp109137554Owttrim GW RNA helicases and abiotic stress Nucleic AcidsRes 200634(11)3220ndash30Park BJ Liu Z Kanno A Kameya T Increased tolerance to salt and water de1047297cit stress in
transgenic lettuce (Lactuca sativa L) by constitutive expression of LEA PlantGrowth Regul 2005a45165ndash71
Park BJ Liu Z Kanno A Kameya T Genetic improvement of Chinese cabbage for salt anddroughttolerance by constitutiveexpression of a B napus LEAgenePlantSci 2005b169553ndash8
Pastori GM Foyer CH Common componentsnetworks and pathways of crosstoleranceto stress The central role of ldquoredoxrdquo and abscisic acid-mediated controls PlantPhysiol 2002129460ndash8
PellegrineschiA ReynoldsM PachecoM Brito RMAlmeraya R Yamaguchi-ShinozakiK et alStress-induced expression in wheat of the Arabidopsis thaliana DREB1Agenedelayswaterstress symptoms under greenhouse conditions Genome 200447493ndash500
Perl A Perl-Treves R Galili S Aviv D Shalgi E Malkin S et al Enhanced oxidative-stressdefense in transgenic potato overexpressing tomato Cu Zn superoxide dismutaseTheor Appl Genet 199385568ndash76
Podlich DW Winkler CR Cooper M Mapping as you go an effective approach formarker-assisted selection of complex traits Crop Sci 2004441560ndash71
182 M Ashraf Biotechnology Advances 28 (2010) 169ndash183
Pratt RC Casey MA Registration of maize germplasm line Oh605 Crop Sci 2006461004ndash5Qin F Kakimoto M Sakuma Y Maruyama K Osakabe Y Tran LSP et al Regulation and
functional analysis of ZmDREB2A in response to drought and heat stress in Zea mays LPlant J 20075054ndash69
Quan R Shang M Zhang H Zhao Y Zhang J Engineering of enhanced glycinebetainesynthesis improves drought tolerance in maize Plant Biotechnol J 20042477ndash86
Quarrie SA Gulli M Calestani C Steed A Marmiroli N Location of a gene regulatingdrought-induced abscisic acid production on the long arm of chromosome 5A of wheat Theor Appl Genet 199489794ndash800
QuickJS Stromberger JA Clayshulte S Clifford B Johnson JJ Peairs FB et al Registrationof lsquoPrairie Redrsquo wheat Crop Sci 2001411362ndash3
RajaramS Prospects andpromise ofwheatbreedingin the21stcentury Euphytica 20011193-15Rajaram S Role of conventional plant breeding and biotechnology in future wheat
production Turk J Agric Forest 200529105ndash11Rami JF Dufour P Trouche G Fliedel G Mestres C Davrieux F et al Quantitative trait
loci for grain quality productivity morphological and agronomical traits insorghum (Sorghum bicolor L Moench) Theor Appl Genet 199897605ndash16
Rassaa N Salahb HBH Latiri K Thermal responses of durum wheat Triticum durum toearly water stress consequence on leaf and 1047298ower development Plant Biol Pathol2008331(5)363ndash71
Reddy LJ Nigam SN Rao RCN Reddy NS Registration of ICGV 87354 peanut germplasmwith drought tolerance and rust resistance Crop Sci 200141274ndash5
Ribaut JM Ragot M Marker-assisted selection to improve drought adaptation in maize thebackcross approach perspectives limitations and alternatives J Exp Bot 200658351ndash60
Robin S Pathan MS Courtois B La1047297tte R Carandang S Lanceras S et al Mappingosmotic adjustment in an advanced back-cross inbred population of rice Theor
Appl Genet 2003107(7)1288ndash96Romero C Belles JM Vaya JL Serrano R Culianez-Macia FA Expression of the yeast
trehalose-6-phosphate synthase gene in transgenic tobacco plants pleiotropicphenotypes include drought tolerance Planta 1997201293ndash7
Ronde JAD Cress WA Krugerd GHJ Strasserd RJ Van Staden J Photosynthetic responseof transgenic soybean plants containing an Arabidopsis P5CR gene during heat anddrought stress J Plant Physiol 20041611211ndash24
Salvi S Tuberosa R To clone or not to clone plant QTLs present and future challengesTrends Plant Sci 200510297ndash304
Sanchez AC Subudhi PK Rosenow DT Nguyen HT Mapping QTLs associated with droughtresistance in sorghum (Sorghum bicolor L Moench) Plant Mol Biol 200248713ndash26
Saranga Y Menz M Jiang CX Wright RJ Yakir D Paterson AH Genomic dissection of genotypetimesenvironment interactions conferring adaptation of cotton to aridconditions Genome Res 2001111988ndash95
Sari-Gorla M Krajewski P Di Fonzo N Villa M Frova C Genetic analysis of droughttolerance in maize by molecular markers II Plant height and 1047298owering Theor ApplGenet 199999289ndash95
Seki M Narusaka M Ishida J Nanjo T Fujita M Oono Y et al Monitoring the expressionpro1047297les of 7000 Arabidopsis genes under drought cold and high-salinity stressesusing a full-length cDNA microarray Plant J 200231279ndash92
Serraj R Sinclair TR Osmolyte accumulation can it really increase crop yield underdrought conditions Plant Cell Environ 200225333ndash41
Serraj R Krishnamurthy L Kashiwagi J Kumar J Chandra S Crouch JH Variation in roottraits of chickpea (Cicer arietinum L) grown under terminal drought Field CropsRes 200488115ndash27
Serraj R Hash CT Rizvi MHS Sharma A Yadav RS Bidinger FR Recent advances in marker-assisted selection for drought tolerance in pearl millet Plant Prod Sci 20058(3)334ndash7
Shen YG Du BX Zhang WK Zhang JS Chen SY AhCMO regulated by stresses in Atriplexhortensis can improve drought tolerance in transgenic tobacco Theor Appl Genet2002105815ndash21
Shinozaki K Yamaguchi-Shinozaki K Molecular responses to dehydration and lowtemperature differences and cross-talk between two stress signaling pathwaysCurr Opin Plant Biol 20003217ndash23
Singh KB Omar M Saxena MC Johansen C Registration of FLIP 87-59C a drought-tolerant chickpea germplasm line Crop Sci 199636(2)1ndash2
Singh SP Teran H Gutierrez JA Registration of SEA 5 and SEA 13 drought tolerant drybean germplasm Crop Sci 200141276ndash7
Singh SP Teran H Lema M Schwartz HF Miklas PN Registration of white moldresistant dry bean germplasm line A 195 J Plant Registr 2007162ndash3
Sivamani E Bahieldin A Wraith JM Improved biomass productivity and water useef 1047297ciency under water de1047297cit conditions in transgenic wheat constitutivelyexpressing the barley HVA1 gene Plant Sci 20001551ndash9
Skovmand B Reynolds MP DeLacy IH Searching genetic resources for physiologicaltraits with potential for increasing yield In Reynolds MP Ortiz-Monasterio IMcNab A editors Application of Physiology in Wheat Breeding 2001 p 17ndash28
Steele K Novel upland rice variety bred using marker-assisted selection and client-oriented breeding released in Jharkhand India Bangor University 2009
Steele KA Price AH Shashidar HE Witcombe JR Marker-assistedselection to introgressrice QTLs controlling root traitsinto an Indianupland rice variety Theor Appl Genet2006112208ndash21
Steele KA Virk DS Kumar R Prasad SC Witcombe JR Field evaluation of upland ricelines selected for QTLs controlling root traits Field Crops Res 2007101180 ndash6
Sunkar R Kapoor A Zhu JK Post transcriptional induction of two CuZn superoxidedismutase genes in Arabidopsis is mediated by down regulation of miR398 andimportant for oxidative stress tolerance Plant Cell 2006182051ndash65
Talame V Sanguineti MC Chiapparino E Bahri H Ben Salem M Forster BP et alIdenti1047297cation of Hordeum spontaneum QTL alleles improving 1047297eld performance of barley grown under rainfed conditions Ann Appl Bot 2004144309ndash20
Tanner NKCordinO BanroquesJ DoereM Linder PThe Q Motif a newlyidenti1047297ed motif in DEAD box helicases may regulate ATP binding and hydrolysis Mol Cell 200311127ndash38
Taramino G Tarchini R Ferrario S Lee M Pe ME Characterization and mapping of simplesequence repeats (SSRs) in Sorghum bicolor Theor Appl Genet 19979566ndash72
Teulat B Monneveux P Wery J Borriegraves C Souyris I Charrier A et al Relationshipsbetween relative water content andgrowth parameters in barleya QTLstudy NewPhytol 199713799-107
Thi Lang N Chi Buu B Fine mapping for drought tolerance in rice ( Oryza sativa L)Omonrice 2008169-15Till BJ Comai L Henikoff S Tillering and ecotillering for crop improvement In
Varshney RK Tuberosa R editors Genomics-assisted crop improvement genomicsapproaches and platforms vol 1 2007 p 333ndash50
Tondelli A Francia E Barabaschi D Aprile A Skinner JS Stockinger EJ et al Mappingregulatory genes as candidates for cold and drought stress tolerance in barleyTheor Appl Genet 2006112445ndash54
Tuberosa R Salvi S Genomics approaches to improve drought tolerance in cropsTrends Plant Sci 200611405ndash12
Valkoun JJ Wheat pre-breeding using wild progenitors Euphytica 200111917ndash23Vashisht AA Tuteja N Stress responsive DEAD-box helicases a new pathway to
engineer plant stress tolerance J Photochem Photobiol B Biol 200684150 ndash60Vendruscolo ECG Schuster I Pileggi M Scapim CA Molinari HBC Marur CJ et al Stress-
induced synthesis of proline confers tolerance to water de1047297cit in transgenic wheat J Plant Physiol 20071641367ndash76
Vienne D Leonardi A Damerval C Zivy M Genetics of proteome variation for QTL characterization application to drought stress responses in maize J Exp Bot 199950303ndash9
Villareal RL Mujeeb-Kazi A Rajaram S Toro ED Morphological variability in somesynthetic hexaploid wheats derived from Triticum turgidum times T tauschii J GenetBreed 1994487-16
Vinh NT Paterson AH Genome mapping and its implication for stress resistance inplants In Ashraf M Harris PJC editors Abiotic stresses plant resistance throughbreeding and molecular approaches 2005
Vinocur B Altman A Recent advances in engineering plant tolerance to abiotic stressachievements and limitations Curr Opin Biotechnol 200516123ndash32
Wang YJ Hao YJ Zhang ZG Chen T Zhang JS Chen SY Isolation of trehalose-6-phosphate phosphatase gene from tobacco and its functional analysis in yeast cells
J Plant Physiol 2005162215ndash23Wu R Garg A Engineering rice plants with trehalose-producing genes improves
tolerance to drought salt and low temperature ISB News Report 2003Xiao B Huang Y Tang N Xiong L Overexpression of LEA gene in rice improves drought
resistance under 1047297eld conditions Theor Appl Genet 200711535ndash46Xinglai P Sangang X Qiannying P Yinhong S Registration of lsquo Jinmai 50rsquo wheat Crop Sci
200646983ndash5Xiong L Lee H Ishitani M Zhu JK Regulation of osmotic stress-responsive gene
expression by the LOS6ABA1 locus in Arabidopsis J Biol Chem 20022778588ndash96Xu GW Magill CW Shertz KF Hart GE A RFLP linkage map of Sorghum bicolor (L)
Moench Theor Appl Genet 199489139ndash45Xu D Duan X Wang B Hong BHo THD Wu RExpression of a late embryogenesis abundant
protein gene HVA1 from barley confers tolerance to water de1047297cit and salt stress intransgenic rice Plant Physiol 1996110249ndash57
Yadav RS Hash CT Bidinger FR Devos KM Howarth CJ Genomic regions associatedwith grain yield and aspects of post-1047298owering drought tolerance in pearl milletacross stress environments and testers background Euphytica 2004136265ndash77
YamadaM MorishitaH Urano K Shiozaki N Yamaguchi-Shinozaki K ShinozakiK et alEffects of free proline accumulation in petunias under drought stress J Exp Bot2005561975ndash81
Yang WJ Rich PJ Axtell JD Wood KV Bonham CC Ejeta G et al Genotypic variation forglycinebetaine in sorghum Crop Sci 200343162ndash9
Yan-Ying QU Ping MU Xue-Qin L Yu-Xiu T Feng W Hong-Liang Z et al QTL mappingand correlations between leaf water potential and drought resistance in rice underupland and lowland environments Acta Agron Sin 200834(2)198ndash206
Ye XDWu XLZhaoH Frehner MNoumlsberger J Potrykus Iet al Altered fructan accumulationin transgenic Lolium multi 1047298orum plants expressing a Bacillus subtilis sacB gene Plant Cell
Rep 200120205ndash
12Zaharieva M Gaulin E Havaux M Acevedo E Monneveux P Drought and heat responses in
the wild wheat relative Aegilops geniculata Roth potential interest for wheatimprovement Crop Sci 2001411321ndash9
Zhang J Zheng HGAartiA Pantuwan G NguyenTT Tripathy JNet alLocatinggenomicregions associated with components of drought resistance in rice comparativemapping within and across species Theor Appl Genet 200110319ndash29
Zhang GH Su Q An LJ Wu S Characterization and expression of a vacuolar Na+H+
antiporter gene from the monocot halophyte Aeluropus littoralis Plant PhysiolBiochem 200846117ndash26
Zhao J QTLs for oil content and their relationships to other agronomic traits in anEuropeantimesChinese oilseed rape population Germany Diss Grorg-Agust Univer-sity of Goettingen 2002
Zhao J Ren W Zhi D Wang L Xia G Arabidopsis DREB1ACBF3 bestowed transgenic tallfescue increased tolerance to drought stress Plant Cell Rep 2007261521ndash8
Zhou W Li Y Zhao BC Ge RC Shen YZ Wang G Huang ZJ Over-expression of TaSTRG gene improves salt and drought tolerance in rice J Plant Physiol 200915(166)1660ndash70
and cost-effective Molecular mapping and analysis of QTL have been
carried out for a number of qualitative and quantitative traits including
stress tolerance which has undoubtedly resulted in a great magnitude
of knowledge and better understanding of the causal genetic phenom-
ena that regulate these traits However limited success has resulted by
using this knowledge to manipulate genes in an effective way for the
improvement of a speci1047297c trait in crops although some crop cultivars
developed through using these tools have performed extremely well
under1047297
eld stressconditions However there are a variety of reasons forthe limited success in terms of achieving enhanced drought tolerance
using the marker-assisted breeding approach For example a major
dif 1047297culty confronting the scientists is the challenge of precise QTL
identi1047297cation In addition a substantial genetictimes environment interac-
tion inconsistent repeatability large number of genes regulating yield
and invalid use of mapping populations have hampered the pursuits
involving mapping of QTL for enhanced drought stress tolerance
Despite these other factors also impede the application of QTL for
genetic improvement of a trait For example due to unfavorable
epistatic interaction it is hard to transfer the effects of a desired allele to
an elite background material (Podlich et al 2004 Collins et al 2008)
Furthermore in some cases QTL from a speci1047297c background do not
show signi1047297cant effects or cease completely in different backgrounds
even under analogous growth conditions (Cho and Hong 2006 Collins
et al 2008) This makes the utilization of QTL more intricate
Transformation of the knowledge acquired from QTL-oriented molec-
ular studies into a well-de1047297ned upshot for the stakeholders is one of the
key challenges confronting the breeders Certainly a multidisciplinary
approach including more speci1047297cally the identi1047297cation of QTL mediated
signal transduction in response to stresses needs to be adopted
Furthermore although QTL cloning procedures are unwieldy they can
lead to a meaningful outcome because a cloned QTL can offer a reliable
markerfor MABas well as furnish an outlinefor thedetection of superior
allelic variants in crop species (Till et al 2007)
Genetic engineering (Transgenic approach) offers a promise whereby
one can expect a substantial improvement in a desired trait within the
shortest stretch of time Engineering genes encoding organic osmolytes
plant growth regulators antioxidants late embryogenesis abundant pro-
teins and transcription factors has resulted into transgenic lines whichhave out-performed under controlled stress conditions In most cases the
claims made by the researchers in terms of performance of transgenic
lines tested under controlled conditions are considerably in1047298ated
However with the exception of a very few notable cases most of the
transgenic lines of differentcrops have been rarely1047297eld-tested Thushow
far these transgenic lines perform under 1047297eld stress conditions is not
known because under natural1047297eld conditions a linecultivar has to face a
multitude of environmental factors other than the drought stress Thus
appraisal of performance of a transgenic line under natural 1047297eld con-
ditionsis mandatory forthe stakeholdersperspective It is also imperative
to note that most of the drought tolerant transgenic lines have been
developed using a single gene transformation which may not be as
productive as if it had been developed using transformation of many
genes Thus transferring a number of prominent genes effectively in-volved in stress tolerance to transgenic plants seems to be a logical
approach Although a large number of genes appear to be involved in
stress tolerance and most of them have been fully characterized the
function of many of them in the mechanism of stress tolerance is yet to be
investigated
References
Abebe T Guenzi AC Martin B Cushman JC Tolerance of mannitol-accumulatingtransgenic wheat to water stress and salinity Plant Physiol 20031311748ndash55
Ashraf M Breeding for salinity tolerance in plants Crit Rev Plant Sci 19941317 ndash42Ashraf M Akram NA Improving salinity tolerance of plants through conventional
breeding and genetic engineering an analytical comparison Biotechnol Adv200927744ndash52 doi101016jbiotechadv200905026
Ashraf M Foolad MR Roles of glycinebetaine and proline in improving plant abioticstress resistance Environ Exp Bot 200759206ndash16
Ashraf M Athar HR Harris PJC Kwon TR Some prospective strategies for improvingcrop salt tolerance Adv Agron 20089745-110
Asins MJ Present and future of quantitative trait locus analysis in plant breeding PlantBreed 2002121281ndash91
Babu RC Nguyen BD Chamarerk V Shanmugasundaram P Chezhian P Jeyaprakash P et alGeneticanalysis of droughtresistance in riceby molecular markers association betweensecondary traits and 1047297eld performance Crop Sci 2003431457ndash69
Babu RC Zhang J Blum A Ho THD Wu R Nguyen HT HVA1 a LEA gene from barleyconfers dehydration tolerance in transgenic rice( Oryzasativa L)via cell membraneprotection Plant Sci 2004166855ndash62
Badawi GH Kawano N Yamauchi Y Over-expression of ascorbate peroxidase in
tobacco chloroplasts enhances the tolerance to salt stress and water de1047297cit PhysiolPlant 2004121231ndash8Badu-Apraku B Yallou CG Registration of striga-resistant and drought tolerant tropical
early maize populations TZE-W Pop DT STR C4 and TZE-Y Pop DT STR C4 J PlantRegistr 20093(1)86ndash90
Badu-Apraku B Menkir A Kling JG Fakorede MAB Registration of 16 striga resistantearly maturing tropical maize inbred lines Crop Sci 2006461410ndash1
Baenziger PS Beecher B Graybosch RA Ibrahim AMH Baltensperger DD Nelson LA et alRegistration of lsquoNEO1643rsquo wheat J Plant Registr 20082(1)36ndash42
Baumlnziger M Setimela PS Hodson D Vivek B Breeding for improved drought tolerance inmaize adapted to southern Africa Proceedings of the 4th International Crop ScienceCongress Brisbane Australia Published on CDROM 2004 26 Sep ndash 1 Oct
Bartels D Sunkar R Drought and salt tolerance in plants Crit Rev Plant Sci 20052423 ndash58Baum M Grandol S Backes G Jahoor A Sabbagh A Ceccarelli S QTLs for agronomic
traits in the Mediterranean environment identi1047297ed in recombinant inbred lines of the cross lsquoArtarsquo H spontaneum 41-1 Theor Appl Genet 20031071215ndash25
Bergman JW Riveland NR Flynn CR Carlson GR Wichman DM Registration of lsquoMorlinrsquo
saf 1047298ower Crop Sci 2001411640Bernier J Kumar A Venuprasad R Spaner D Atlin G A large-effect QTL for
grain yield under reproductive-stage drought stress in upland rice Crop Sci200747507ndash18
BernierJ Kumar A SerrajR SpanerD Atlin G Review breeding uplandrice fordroughtresistance J Sci Food Agric 200888927ndash39
Bernier J Serraj R Kumar A Venuprasad R Impa S Gowdaa RPV et al The large-effectdrought-resistance QTL qtl121 increases water uptake in upland rice Field CropsRes 200911039ndash46
Bhatnagar-Mathur P ReddyDS Lavanya M Yamaguchi-Shinozaki K Sharma KK Stress-inducible expression of Arabidopsis thaliana DREB1A in transgenic peanut ( Arachishypogaea L) increases transpiration ef 1047297ciency under water-limiting conditionsPlant Cell Rep 2007262071ndash82
Bhatnagar-Mathur P Devi MJ Vadez V Sharma KK Differential antioxidative responses intransgenic peanut bear no relationship to their superior transpiration ef 1047297ciency underdrought stress J Plant Physiol 2009166(11)1207ndash17 doi101016jjplph200901001
Bidinger FR Serraj R Rizvi SMH Howarth C Yadav RS Hash CT Field evaluation of droughttolerance QTL effects on phenotype and adaptation in pearl millet [ Pennisetum glaucum(L) R Br] topcross hybrids Field Crops Res 200594(1)14ndash32
Bidinger FR Nepolean T Hash CT Yadav RS Howarth CJ Identi1047297cation of QTLs for grainyield of pearl millet (Pennisetum glaucum (L) R Br) in environments with variablemoisture during grain 1047297lling Crop Sci 200747969ndash80
Bowers JE Abbey C Anderson S Chang C Draye X A high-density geneticrecombination map of sequence-tagged sites for Sorghum as a framework forcomparative structural and evolutionary genomics of tropical grains and grassesGenetics 2003165367ndash86
Brick MA Ogg JB Singh SP Schwartz HF Johnson JJ Pastor-Corrales MA Registration of drought-tolerant rust-resistant high-yielding pinto bean germplasm lineCO46348 J Plant Registr 20082(2)120ndash4
Browne J Tunnacliffe A Burnell A Anhydrobiosis-plant desiccation gene found in anematode Nature 200241638
Busk PK Pages M Regulation of abscisic acid induced transcription Plant Mol Biol199837425ndash35
Carena MJ Wanner DW Development of genetically broad-based inbred lines of maizefor early-maturing (70-80RM) hybrids J Plant Registr 20093107ndash11
Cash SDBrucknerPL Wichman DMKephart KD Berg JEBoynerR et alRegistration of Willow Creek forage wheat J Plant Registr 20093(2)185ndash90
Cattivelli L Rizza F Badeck FW Mazzucotelli E Francia AMEM Mare AT et al Droughttolerance improvement in crop plants an integrated view from breeding to genomics
Field Crops Res 20081051-14Chen M Wang QY Cheng XG Xu ZS Li LC Ye XG et al GmDREB2 a soybean DRE-
binding transcription factor conferred drought and high-salt tolerance intransgenic plants Biochem Biophys Res Commun 2007353299ndash305
Cheng Z Targolli J Huang X Wu R Wheat LEA genes PMA80 and PMA1959 enhancedehydration tolerance of transgenic rice (Oryza sativa L) Mol Breed 20021071ndash82
ChoEK Hong ChB Over-expression of tobacco NtHSP70-1 contributes to drought-stresstolerance in plants Plant Cell Reports 200625349ndash58
Close TJ Dehydrins a commonality in the response of plants to dehydration and lowtemperature Physiol Plant 1997100291ndash6
Collins NC Tardieu F Tuberosa R Quantitative trait loci and crop performance underabiotic stress where do we stand Plant Physiol 2008147469ndash86
Concept Note Combining breeding and biotechnology to develop water ef 1047297cient maizefor Africa (WEMA) Afr Agric Technol Foundation
Courtois BShen L Petalcorin WCarandang S MauleonR LiZ Locating QTLs controllingconstitutive root traits in the rice population IAC 165-Co39 Euphytica 2003134335ndash45
Da-hong L Hui L Yang YL Ping-ping Z Jian-sheng L Down-regulated expression of RACK1geneby RNAinterference enhancesdroughttolerancein rice RiceSci 200916(1)14ndash20
Dalal M Tayal D Chinnusamy V Bansala KC Abiotic stress and ABA-inducible group 4 LEAfrom Brassicanapus plays a key rolein saltand droughttolerance J Biotechnol 2009139137ndash45
Ding Z Li S An X Liu X Qin H Wang D Transgenic expression of MYB15 confers enhancedsensitivity to abscisic acid and improved drought tolerance in Arabidopsis thaliana
J Genet Genomics 20093617ndash29Dubouzet JG Sakuma Y Ito Y Kasuga M Dubouzet EG Miura S et al OsDREB genes in
rice Oryza sativa L encode transcription activators that function in drought high-salt- and cold-responsive gene expression Plant J 200333751ndash63
El-Far IA Allan AY Responses of some wheat cultivars to sowing methods and droughtat different stages of growth Assuit J Agric Sci 199526(1)267ndash77
Eltayeb AE Kawano N Badawi GH Kaminaka H Sanekata T Shibahara T et alOverexpression of monodehydroascorbate reductase in transgenic tobacco confersenhanced tolerance to ozonesalt and polyethyleneglycol stresses Planta 2007225(5)1255ndash64
Falconer DS Introduction to quantitative genetics London New York Longman 1989Feng-ling FU Zhi-Lei F Shi-bing G Shu-feng Z Wan-chen L Evaluation and quantitative
inheritanceof several drought-relativetraits in maize Agric SciChina 20087(3)280ndash90Finkelstein R Gampala S Rock C Abscisic acid signaling in seeds and seedlings Plant
Cell 20021415ndash45Flowers TJ Improving crop salt tolerance J Exp Bot 200455307ndash19Fujita Y Fujita M Satoh R Maruyama K Parvez MM Seki M et al AREB1 is a transcription
Giraudat J Parcy F Bertauche N Gosti F Leung J Morris PC et al Current advances inabscisic acid action and signalling Plant Mol Biol 1994261557ndash77
Gorbalenya AE Koonin EV Helicases amino acid sequence comparisons and structurendashfunction relationships Curr Opin Struct Biol 19933419ndash29
Gosal SS Wani SH Kang MS Biotechnology and drought tolerance J Crop Improvement20092319ndash54
Gubis J Vaňkovaacute R Červenaacute V Draguacuteňovaacute M Hudcovicovaacute M Lichtnerovaacute H et alTransformedtobacco plants with increased toleranceto droughtSouthAfr J Bot200773505ndash11
GuoO Zhang J GaoQ Xing SLi F Wang WDrought tolerancethrough over-expressionof mono ubiquitin in transgenic tobacco J Plant Physiol 20081651745 ndash55
Haley SD Johnson JJ Peairs FB Quick JS Stromberger JA Clayshulte SR et al Registration of lsquoRipperrsquo wheat J Plant Registr 200711ndash6
Han SE Park SR Kwon HB Yi BY Lee GB Byun MO Genetic engineering of drought-resistant tobacco plants by introducingthe trehalose phosphorylase (TP) gene fromPleurotus sajor-caju Plant Cell Tissue Organ Cult 200582151ndash8
Harris K Klein R Mullet J Sorghum stay-green QTL individually reduces post-1047298oweringdrought-induced leaf senescence J Exp Bot 200758327ndash38
Hong BS Zong-Suo L Ming-An S LEA proteins in higher plants structure functiongene expression and regulation Colloids Surf B Biointerf 200545131ndash5
Howarth CJ Yadav RS Successful marker assisted selection for drought tolerance anddisease resistance in pearl milletIGER Innovations 2002
Humphreys MO Humphreys MW Breeding for stress resistance general principles InAshraf M Harris PJC editors Abiotic stresses plant resistance through breedingand molecular approaches 2005 p 19ndash46
Ingram J Bartels D The molecular basis of dehydration tolerance in plants Ann RevPlant Physiol Plant Mol Biol 199647377ndash403
Jaglo-Ottosen KR Gilmour SJ Zarka DG Schabenberger O Thomashow MF ArabidopsisCBF1 overexpression induces COR genes and enhances freezing tolerance Science1998280104ndash6
Jang IC Oh SJ Seo JS Choi WB Song SI Kim CH et al Expression of a bifunctional fusion of the Escherichia coli genes for trehalose-6-phosphate synthase and trehalose-6-phosphate phosphatase in transgenic rice plants increases trehalose accumulationandabiotic stresstolerancewithoutstuntinggrowth Plant Physiol2003131516ndash24
Jeanneau M Gerentes D Foueillassar X Zivy M Vidal J Toppan A et al Improvement of drought tolerance in maize towards the functional validation of the Zm-Asr1 gene andincrease of water use ef 1047297ciency by over-expressing C4-PEPC Biochimie 2002841127ndash35
Jenson KB Palazzo AJ Waldron BL Bushman BS Registration of lsquoFirstStrikersquo Slenderwheatgrass J Plant Registr 2007124ndash5
Joung-youn K Hyung-in C Min-young I Soo-young K Arabidopsis basic leucine zipperproteins that mediate stress-responsive abscisic acid signaling Plant Cell 20024343ndash57
Juskiw PE Helm JH Oro M Nyachiro JM Salmon DF Registration of lsquoBentleyrsquo barley
J Plant Registr 20093(2)119ndash23Kalamaki MS Alexandrou D Lazari D Merkouropoulos G Fotopoulos V Pateraki I et al
Over-expression of a tomato N-acetyl-L -glutamate synthase gene (SlNAGS1) in Arabidopsis thaliana results in high ornithine levels and increased tolerance in saltand drought stresses J Exp Bot 200960(6)1859ndash71
Kamoshita A Babu CR Boopathi NM Fukai S Phenotypic and genotypic analysis of drought-resistance traits for development of rice cultivars adapted to rainfedenvironments Field Crops Res 2008109(103)1-23 doi101016jfcr200806010
KarakasB Ozias-AkinsP Stushnoff C SuefferheldM Rieger M Salinityand drought toleranceof mannitol-accumulating transgenic tobacco Plant Cell Environ 199720609ndash16
Karim S Aronsson H Ericson H Pirhonen M Leyman B Welin B et al Improved droughttolerance without undesired side effects in transgenic plants producing trehalose PlantMol Biol 200764371ndash86
Kasuga M Liu Q Miura S Yamaguchi-Shinozaki K Shinozaki K Improving plant droughtsaltand freezing tolerance by gene transfer of a single stress-inducible transcription factorNat Biotechnol 199917287ndash91
Kindiger M Gaub H Hasegawac M Katsurab Y Ueyamad K Gotob S et al Registrationof lsquoNanryorsquo tall fescue Crop Sci 2006461815ndash6
Kong I Dong J HartGE Characteristics linkage mappositionsand allelic differentiationof Sorghum bicolour (L) Moench by DNA simple-sequence repeats (SSRs) TheorAppl Genet 2000101438ndash48
Kumar R Venuprasad R Atlin GN Genetic analysis of rainfed lowland rice droughttolerance under naturally-occurring stress in eastern India heritability and QTL effects Field Crops Res 200710342ndash52
La1047297tte HR Price AH Courtois B Yield response to water de1047297cit in an upland ricemapping population associations among traits and genetic markers Theor ApplGenet 20041091237ndash46
Lal S Gulyani V Khurana P Overexpressionof HVA1 gene frombarley generates tolerance tosalinity and water stress in transgenic mulberry (Morus indica) Transgenic Res 200817
(4)651ndash
63Lanceras J Pantuwan G Jongdee B Toojinda T Quantitative trait loci associated withdrought tolerance at reproductive stage in rice Plant Physiol 2004135384ndash99
Levi A Ovnat L Paterson AH Saranga Y Photosynthesis of cotton near-isogenic linesintrogressed with QTLs for productivity and drought related traits Plant Sci 2009a17788ndash96
Levi A Paterson AH Barak V Yakir D Wang B Chee PW et al Field evaluation of cottonnear-isogenic lines introgressed with QTLs for productivity and drought relatedtraits Mol Breed 2009b23179ndash95
Liu Q Kasuga M Sakuma Y Abe H Miura S Yamaguchi- Shinozaki K et al Twotranscription factors DREB1 and DREB2 with an EREBPAP2 DNA binding domainseparate two cellular signal transduction pathways in drought-and low-temper-ature-responsive gene expression respectively in Arabidopsis Plant Cell 1998101391ndash406
Liu X Hua X Guo J Qi D Wang L Liu Z et al Enhanced tolerance to drought stress intransgenic tobacco plants overexpressing Biotechnol Lett 2008301275ndash80
Liu X Wanga Z Wanga L Wua R Phillips J Deng X LEA 4 group genes from theresurrection plant Boea hygrometrica confer dehydration tolerance in transgenictobacco Plant Sci 200917690ndash8
Luchi S Kobayashi M Taji T Naramoto M Seki M Kato T et al Regulation of droughttolerance by gene manipulation of 9-cis-epoxycarotenoid dioxygenase a keyenzyme in abscisic acid biosynthesis in Arabidopsis Plant J 200127325ndash33
Luo Y Liu YB Dong YX Gao XQ Zhang YS Expression of a putative alfalfa helicase increasestolerance to abioticstress in Arabidopsis by enhancing the capacities for ROS scavengingand osmotic adjustment J Plant Physiol 2009166385ndash94
MacLean JLDaweDC Hardy B Hettel GPRiceAlmanac sourcebook for themostimportanteconomic activity on Earth3rd ed Wallingford England CABI Publishing 2002
Mansour MMF Nitrogen containing compounds and adaptation of plants to salinitystress Biol Plant 200043491ndash500
McKersie BD Bowley SR Harjanto E Leprince O Water-de1047297cit tolerance and 1047297eldperformance of transgenic alfalfa overexpressing superoxide dismutase Plant Physiol19961111177ndash81
McKersie BD Murnaghan J Bowley SR Manipulating freezing tolerance in transgenicplants Acta Physiol Plant 199719485ndash95
Miranda JA Avonce N Suaacuterez R Thevelein JM Dijck PV Iturriaga G A bifunctional TPS-TPP enzyme from yeast confers tolerance to multiple and extreme abiotic-stressconditions in transgenic Arabidopsis Planta 2007226(6)1411ndash21
Mohammadi M Taleei A Zeinali H Naghavi MR Ceccarelli S Grando Baum M QTL analysis for phenologic traits in doubled haploid population of barley Int J AgricBiol 20057(5)820ndash3
Mohanty A Kathuria H Ferjani A Sakamoto A Mohanty P Murata N et al Transgenicsof an elite indica rice variety Pusa Basmati 1 harbouring the codA gene are highlytolerant to salt stress Theor Appl Genet 200210651ndash7
Noaman MM El Sayad AA Asaad FA El Sherbini AM El Bawab AO El Moselhi MA et alRegistration of lsquoGiza 126rsquo barley Crop Sci 199535(6)1710
Noaman MM Ahmed IA El-Sayed AA Abo-El-Enin RA El-Gamal AS El-Sherbiny AM et alRegistration of lsquoGiza 2000rsquo drought-tolerant six-rowed barley for rainfed and newreclaimed areas in Egypt Crop Sci 200747440
Obert DE Evans CP Wesenberg DM Windes JM Erickson CA Jackson EW et alRegistration of lsquoLenetahrsquo spring barley J Plant Registr 20082(2)85ndash7
Oh SJ Jeong JS Kim EH Yi NR Yi SI Jang IC et al Matrix attachment region fromthe chicken lysozyme locus reduces variability in transgene expression andconfers copy number-dependence in transgenic rice plants Plant Cell Rep20054145ndash54
Oh SJ Kim YS Kwon C Park HK Jeong JS Kim JK Overexpression of the transcriptionfactor AP37 in rice improves grain yield under drought conditions Plant Physiol
20092191ndash
200 doi101104pp109137554Owttrim GW RNA helicases and abiotic stress Nucleic AcidsRes 200634(11)3220ndash30Park BJ Liu Z Kanno A Kameya T Increased tolerance to salt and water de1047297cit stress in
transgenic lettuce (Lactuca sativa L) by constitutive expression of LEA PlantGrowth Regul 2005a45165ndash71
Park BJ Liu Z Kanno A Kameya T Genetic improvement of Chinese cabbage for salt anddroughttolerance by constitutiveexpression of a B napus LEAgenePlantSci 2005b169553ndash8
Pastori GM Foyer CH Common componentsnetworks and pathways of crosstoleranceto stress The central role of ldquoredoxrdquo and abscisic acid-mediated controls PlantPhysiol 2002129460ndash8
PellegrineschiA ReynoldsM PachecoM Brito RMAlmeraya R Yamaguchi-ShinozakiK et alStress-induced expression in wheat of the Arabidopsis thaliana DREB1Agenedelayswaterstress symptoms under greenhouse conditions Genome 200447493ndash500
Perl A Perl-Treves R Galili S Aviv D Shalgi E Malkin S et al Enhanced oxidative-stressdefense in transgenic potato overexpressing tomato Cu Zn superoxide dismutaseTheor Appl Genet 199385568ndash76
Podlich DW Winkler CR Cooper M Mapping as you go an effective approach formarker-assisted selection of complex traits Crop Sci 2004441560ndash71
182 M Ashraf Biotechnology Advances 28 (2010) 169ndash183
Pratt RC Casey MA Registration of maize germplasm line Oh605 Crop Sci 2006461004ndash5Qin F Kakimoto M Sakuma Y Maruyama K Osakabe Y Tran LSP et al Regulation and
functional analysis of ZmDREB2A in response to drought and heat stress in Zea mays LPlant J 20075054ndash69
Quan R Shang M Zhang H Zhao Y Zhang J Engineering of enhanced glycinebetainesynthesis improves drought tolerance in maize Plant Biotechnol J 20042477ndash86
Quarrie SA Gulli M Calestani C Steed A Marmiroli N Location of a gene regulatingdrought-induced abscisic acid production on the long arm of chromosome 5A of wheat Theor Appl Genet 199489794ndash800
QuickJS Stromberger JA Clayshulte S Clifford B Johnson JJ Peairs FB et al Registrationof lsquoPrairie Redrsquo wheat Crop Sci 2001411362ndash3
RajaramS Prospects andpromise ofwheatbreedingin the21stcentury Euphytica 20011193-15Rajaram S Role of conventional plant breeding and biotechnology in future wheat
production Turk J Agric Forest 200529105ndash11Rami JF Dufour P Trouche G Fliedel G Mestres C Davrieux F et al Quantitative trait
loci for grain quality productivity morphological and agronomical traits insorghum (Sorghum bicolor L Moench) Theor Appl Genet 199897605ndash16
Rassaa N Salahb HBH Latiri K Thermal responses of durum wheat Triticum durum toearly water stress consequence on leaf and 1047298ower development Plant Biol Pathol2008331(5)363ndash71
Reddy LJ Nigam SN Rao RCN Reddy NS Registration of ICGV 87354 peanut germplasmwith drought tolerance and rust resistance Crop Sci 200141274ndash5
Ribaut JM Ragot M Marker-assisted selection to improve drought adaptation in maize thebackcross approach perspectives limitations and alternatives J Exp Bot 200658351ndash60
Robin S Pathan MS Courtois B La1047297tte R Carandang S Lanceras S et al Mappingosmotic adjustment in an advanced back-cross inbred population of rice Theor
Appl Genet 2003107(7)1288ndash96Romero C Belles JM Vaya JL Serrano R Culianez-Macia FA Expression of the yeast
trehalose-6-phosphate synthase gene in transgenic tobacco plants pleiotropicphenotypes include drought tolerance Planta 1997201293ndash7
Ronde JAD Cress WA Krugerd GHJ Strasserd RJ Van Staden J Photosynthetic responseof transgenic soybean plants containing an Arabidopsis P5CR gene during heat anddrought stress J Plant Physiol 20041611211ndash24
Salvi S Tuberosa R To clone or not to clone plant QTLs present and future challengesTrends Plant Sci 200510297ndash304
Sanchez AC Subudhi PK Rosenow DT Nguyen HT Mapping QTLs associated with droughtresistance in sorghum (Sorghum bicolor L Moench) Plant Mol Biol 200248713ndash26
Saranga Y Menz M Jiang CX Wright RJ Yakir D Paterson AH Genomic dissection of genotypetimesenvironment interactions conferring adaptation of cotton to aridconditions Genome Res 2001111988ndash95
Sari-Gorla M Krajewski P Di Fonzo N Villa M Frova C Genetic analysis of droughttolerance in maize by molecular markers II Plant height and 1047298owering Theor ApplGenet 199999289ndash95
Seki M Narusaka M Ishida J Nanjo T Fujita M Oono Y et al Monitoring the expressionpro1047297les of 7000 Arabidopsis genes under drought cold and high-salinity stressesusing a full-length cDNA microarray Plant J 200231279ndash92
Serraj R Sinclair TR Osmolyte accumulation can it really increase crop yield underdrought conditions Plant Cell Environ 200225333ndash41
Serraj R Krishnamurthy L Kashiwagi J Kumar J Chandra S Crouch JH Variation in roottraits of chickpea (Cicer arietinum L) grown under terminal drought Field CropsRes 200488115ndash27
Serraj R Hash CT Rizvi MHS Sharma A Yadav RS Bidinger FR Recent advances in marker-assisted selection for drought tolerance in pearl millet Plant Prod Sci 20058(3)334ndash7
Shen YG Du BX Zhang WK Zhang JS Chen SY AhCMO regulated by stresses in Atriplexhortensis can improve drought tolerance in transgenic tobacco Theor Appl Genet2002105815ndash21
Shinozaki K Yamaguchi-Shinozaki K Molecular responses to dehydration and lowtemperature differences and cross-talk between two stress signaling pathwaysCurr Opin Plant Biol 20003217ndash23
Singh KB Omar M Saxena MC Johansen C Registration of FLIP 87-59C a drought-tolerant chickpea germplasm line Crop Sci 199636(2)1ndash2
Singh SP Teran H Gutierrez JA Registration of SEA 5 and SEA 13 drought tolerant drybean germplasm Crop Sci 200141276ndash7
Singh SP Teran H Lema M Schwartz HF Miklas PN Registration of white moldresistant dry bean germplasm line A 195 J Plant Registr 2007162ndash3
Sivamani E Bahieldin A Wraith JM Improved biomass productivity and water useef 1047297ciency under water de1047297cit conditions in transgenic wheat constitutivelyexpressing the barley HVA1 gene Plant Sci 20001551ndash9
Skovmand B Reynolds MP DeLacy IH Searching genetic resources for physiologicaltraits with potential for increasing yield In Reynolds MP Ortiz-Monasterio IMcNab A editors Application of Physiology in Wheat Breeding 2001 p 17ndash28
Steele K Novel upland rice variety bred using marker-assisted selection and client-oriented breeding released in Jharkhand India Bangor University 2009
Steele KA Price AH Shashidar HE Witcombe JR Marker-assistedselection to introgressrice QTLs controlling root traitsinto an Indianupland rice variety Theor Appl Genet2006112208ndash21
Steele KA Virk DS Kumar R Prasad SC Witcombe JR Field evaluation of upland ricelines selected for QTLs controlling root traits Field Crops Res 2007101180 ndash6
Sunkar R Kapoor A Zhu JK Post transcriptional induction of two CuZn superoxidedismutase genes in Arabidopsis is mediated by down regulation of miR398 andimportant for oxidative stress tolerance Plant Cell 2006182051ndash65
Talame V Sanguineti MC Chiapparino E Bahri H Ben Salem M Forster BP et alIdenti1047297cation of Hordeum spontaneum QTL alleles improving 1047297eld performance of barley grown under rainfed conditions Ann Appl Bot 2004144309ndash20
Tanner NKCordinO BanroquesJ DoereM Linder PThe Q Motif a newlyidenti1047297ed motif in DEAD box helicases may regulate ATP binding and hydrolysis Mol Cell 200311127ndash38
Taramino G Tarchini R Ferrario S Lee M Pe ME Characterization and mapping of simplesequence repeats (SSRs) in Sorghum bicolor Theor Appl Genet 19979566ndash72
Teulat B Monneveux P Wery J Borriegraves C Souyris I Charrier A et al Relationshipsbetween relative water content andgrowth parameters in barleya QTLstudy NewPhytol 199713799-107
Thi Lang N Chi Buu B Fine mapping for drought tolerance in rice ( Oryza sativa L)Omonrice 2008169-15Till BJ Comai L Henikoff S Tillering and ecotillering for crop improvement In
Varshney RK Tuberosa R editors Genomics-assisted crop improvement genomicsapproaches and platforms vol 1 2007 p 333ndash50
Tondelli A Francia E Barabaschi D Aprile A Skinner JS Stockinger EJ et al Mappingregulatory genes as candidates for cold and drought stress tolerance in barleyTheor Appl Genet 2006112445ndash54
Tuberosa R Salvi S Genomics approaches to improve drought tolerance in cropsTrends Plant Sci 200611405ndash12
Valkoun JJ Wheat pre-breeding using wild progenitors Euphytica 200111917ndash23Vashisht AA Tuteja N Stress responsive DEAD-box helicases a new pathway to
engineer plant stress tolerance J Photochem Photobiol B Biol 200684150 ndash60Vendruscolo ECG Schuster I Pileggi M Scapim CA Molinari HBC Marur CJ et al Stress-
induced synthesis of proline confers tolerance to water de1047297cit in transgenic wheat J Plant Physiol 20071641367ndash76
Vienne D Leonardi A Damerval C Zivy M Genetics of proteome variation for QTL characterization application to drought stress responses in maize J Exp Bot 199950303ndash9
Villareal RL Mujeeb-Kazi A Rajaram S Toro ED Morphological variability in somesynthetic hexaploid wheats derived from Triticum turgidum times T tauschii J GenetBreed 1994487-16
Vinh NT Paterson AH Genome mapping and its implication for stress resistance inplants In Ashraf M Harris PJC editors Abiotic stresses plant resistance throughbreeding and molecular approaches 2005
Vinocur B Altman A Recent advances in engineering plant tolerance to abiotic stressachievements and limitations Curr Opin Biotechnol 200516123ndash32
Wang YJ Hao YJ Zhang ZG Chen T Zhang JS Chen SY Isolation of trehalose-6-phosphate phosphatase gene from tobacco and its functional analysis in yeast cells
J Plant Physiol 2005162215ndash23Wu R Garg A Engineering rice plants with trehalose-producing genes improves
tolerance to drought salt and low temperature ISB News Report 2003Xiao B Huang Y Tang N Xiong L Overexpression of LEA gene in rice improves drought
resistance under 1047297eld conditions Theor Appl Genet 200711535ndash46Xinglai P Sangang X Qiannying P Yinhong S Registration of lsquo Jinmai 50rsquo wheat Crop Sci
200646983ndash5Xiong L Lee H Ishitani M Zhu JK Regulation of osmotic stress-responsive gene
expression by the LOS6ABA1 locus in Arabidopsis J Biol Chem 20022778588ndash96Xu GW Magill CW Shertz KF Hart GE A RFLP linkage map of Sorghum bicolor (L)
Moench Theor Appl Genet 199489139ndash45Xu D Duan X Wang B Hong BHo THD Wu RExpression of a late embryogenesis abundant
protein gene HVA1 from barley confers tolerance to water de1047297cit and salt stress intransgenic rice Plant Physiol 1996110249ndash57
Yadav RS Hash CT Bidinger FR Devos KM Howarth CJ Genomic regions associatedwith grain yield and aspects of post-1047298owering drought tolerance in pearl milletacross stress environments and testers background Euphytica 2004136265ndash77
YamadaM MorishitaH Urano K Shiozaki N Yamaguchi-Shinozaki K ShinozakiK et alEffects of free proline accumulation in petunias under drought stress J Exp Bot2005561975ndash81
Yang WJ Rich PJ Axtell JD Wood KV Bonham CC Ejeta G et al Genotypic variation forglycinebetaine in sorghum Crop Sci 200343162ndash9
Yan-Ying QU Ping MU Xue-Qin L Yu-Xiu T Feng W Hong-Liang Z et al QTL mappingand correlations between leaf water potential and drought resistance in rice underupland and lowland environments Acta Agron Sin 200834(2)198ndash206
Ye XDWu XLZhaoH Frehner MNoumlsberger J Potrykus Iet al Altered fructan accumulationin transgenic Lolium multi 1047298orum plants expressing a Bacillus subtilis sacB gene Plant Cell
Rep 200120205ndash
12Zaharieva M Gaulin E Havaux M Acevedo E Monneveux P Drought and heat responses in
the wild wheat relative Aegilops geniculata Roth potential interest for wheatimprovement Crop Sci 2001411321ndash9
Zhang J Zheng HGAartiA Pantuwan G NguyenTT Tripathy JNet alLocatinggenomicregions associated with components of drought resistance in rice comparativemapping within and across species Theor Appl Genet 200110319ndash29
Zhang GH Su Q An LJ Wu S Characterization and expression of a vacuolar Na+H+
antiporter gene from the monocot halophyte Aeluropus littoralis Plant PhysiolBiochem 200846117ndash26
Zhao J QTLs for oil content and their relationships to other agronomic traits in anEuropeantimesChinese oilseed rape population Germany Diss Grorg-Agust Univer-sity of Goettingen 2002
Zhao J Ren W Zhi D Wang L Xia G Arabidopsis DREB1ACBF3 bestowed transgenic tallfescue increased tolerance to drought stress Plant Cell Rep 2007261521ndash8
Zhou W Li Y Zhao BC Ge RC Shen YZ Wang G Huang ZJ Over-expression of TaSTRG gene improves salt and drought tolerance in rice J Plant Physiol 200915(166)1660ndash70
and cost-effective Molecular mapping and analysis of QTL have been
carried out for a number of qualitative and quantitative traits including
stress tolerance which has undoubtedly resulted in a great magnitude
of knowledge and better understanding of the causal genetic phenom-
ena that regulate these traits However limited success has resulted by
using this knowledge to manipulate genes in an effective way for the
improvement of a speci1047297c trait in crops although some crop cultivars
developed through using these tools have performed extremely well
under1047297
eld stressconditions However there are a variety of reasons forthe limited success in terms of achieving enhanced drought tolerance
using the marker-assisted breeding approach For example a major
dif 1047297culty confronting the scientists is the challenge of precise QTL
identi1047297cation In addition a substantial genetictimes environment interac-
tion inconsistent repeatability large number of genes regulating yield
and invalid use of mapping populations have hampered the pursuits
involving mapping of QTL for enhanced drought stress tolerance
Despite these other factors also impede the application of QTL for
genetic improvement of a trait For example due to unfavorable
epistatic interaction it is hard to transfer the effects of a desired allele to
an elite background material (Podlich et al 2004 Collins et al 2008)
Furthermore in some cases QTL from a speci1047297c background do not
show signi1047297cant effects or cease completely in different backgrounds
even under analogous growth conditions (Cho and Hong 2006 Collins
et al 2008) This makes the utilization of QTL more intricate
Transformation of the knowledge acquired from QTL-oriented molec-
ular studies into a well-de1047297ned upshot for the stakeholders is one of the
key challenges confronting the breeders Certainly a multidisciplinary
approach including more speci1047297cally the identi1047297cation of QTL mediated
signal transduction in response to stresses needs to be adopted
Furthermore although QTL cloning procedures are unwieldy they can
lead to a meaningful outcome because a cloned QTL can offer a reliable
markerfor MABas well as furnish an outlinefor thedetection of superior
allelic variants in crop species (Till et al 2007)
Genetic engineering (Transgenic approach) offers a promise whereby
one can expect a substantial improvement in a desired trait within the
shortest stretch of time Engineering genes encoding organic osmolytes
plant growth regulators antioxidants late embryogenesis abundant pro-
teins and transcription factors has resulted into transgenic lines whichhave out-performed under controlled stress conditions In most cases the
claims made by the researchers in terms of performance of transgenic
lines tested under controlled conditions are considerably in1047298ated
However with the exception of a very few notable cases most of the
transgenic lines of differentcrops have been rarely1047297eld-tested Thushow
far these transgenic lines perform under 1047297eld stress conditions is not
known because under natural1047297eld conditions a linecultivar has to face a
multitude of environmental factors other than the drought stress Thus
appraisal of performance of a transgenic line under natural 1047297eld con-
ditionsis mandatory forthe stakeholdersperspective It is also imperative
to note that most of the drought tolerant transgenic lines have been
developed using a single gene transformation which may not be as
productive as if it had been developed using transformation of many
genes Thus transferring a number of prominent genes effectively in-volved in stress tolerance to transgenic plants seems to be a logical
approach Although a large number of genes appear to be involved in
stress tolerance and most of them have been fully characterized the
function of many of them in the mechanism of stress tolerance is yet to be
investigated
References
Abebe T Guenzi AC Martin B Cushman JC Tolerance of mannitol-accumulatingtransgenic wheat to water stress and salinity Plant Physiol 20031311748ndash55
Ashraf M Breeding for salinity tolerance in plants Crit Rev Plant Sci 19941317 ndash42Ashraf M Akram NA Improving salinity tolerance of plants through conventional
breeding and genetic engineering an analytical comparison Biotechnol Adv200927744ndash52 doi101016jbiotechadv200905026
Ashraf M Foolad MR Roles of glycinebetaine and proline in improving plant abioticstress resistance Environ Exp Bot 200759206ndash16
Ashraf M Athar HR Harris PJC Kwon TR Some prospective strategies for improvingcrop salt tolerance Adv Agron 20089745-110
Asins MJ Present and future of quantitative trait locus analysis in plant breeding PlantBreed 2002121281ndash91
Babu RC Nguyen BD Chamarerk V Shanmugasundaram P Chezhian P Jeyaprakash P et alGeneticanalysis of droughtresistance in riceby molecular markers association betweensecondary traits and 1047297eld performance Crop Sci 2003431457ndash69
Babu RC Zhang J Blum A Ho THD Wu R Nguyen HT HVA1 a LEA gene from barleyconfers dehydration tolerance in transgenic rice( Oryzasativa L)via cell membraneprotection Plant Sci 2004166855ndash62
Badawi GH Kawano N Yamauchi Y Over-expression of ascorbate peroxidase in
tobacco chloroplasts enhances the tolerance to salt stress and water de1047297cit PhysiolPlant 2004121231ndash8Badu-Apraku B Yallou CG Registration of striga-resistant and drought tolerant tropical
early maize populations TZE-W Pop DT STR C4 and TZE-Y Pop DT STR C4 J PlantRegistr 20093(1)86ndash90
Badu-Apraku B Menkir A Kling JG Fakorede MAB Registration of 16 striga resistantearly maturing tropical maize inbred lines Crop Sci 2006461410ndash1
Baenziger PS Beecher B Graybosch RA Ibrahim AMH Baltensperger DD Nelson LA et alRegistration of lsquoNEO1643rsquo wheat J Plant Registr 20082(1)36ndash42
Baumlnziger M Setimela PS Hodson D Vivek B Breeding for improved drought tolerance inmaize adapted to southern Africa Proceedings of the 4th International Crop ScienceCongress Brisbane Australia Published on CDROM 2004 26 Sep ndash 1 Oct
Bartels D Sunkar R Drought and salt tolerance in plants Crit Rev Plant Sci 20052423 ndash58Baum M Grandol S Backes G Jahoor A Sabbagh A Ceccarelli S QTLs for agronomic
traits in the Mediterranean environment identi1047297ed in recombinant inbred lines of the cross lsquoArtarsquo H spontaneum 41-1 Theor Appl Genet 20031071215ndash25
Bergman JW Riveland NR Flynn CR Carlson GR Wichman DM Registration of lsquoMorlinrsquo
saf 1047298ower Crop Sci 2001411640Bernier J Kumar A Venuprasad R Spaner D Atlin G A large-effect QTL for
grain yield under reproductive-stage drought stress in upland rice Crop Sci200747507ndash18
BernierJ Kumar A SerrajR SpanerD Atlin G Review breeding uplandrice fordroughtresistance J Sci Food Agric 200888927ndash39
Bernier J Serraj R Kumar A Venuprasad R Impa S Gowdaa RPV et al The large-effectdrought-resistance QTL qtl121 increases water uptake in upland rice Field CropsRes 200911039ndash46
Bhatnagar-Mathur P ReddyDS Lavanya M Yamaguchi-Shinozaki K Sharma KK Stress-inducible expression of Arabidopsis thaliana DREB1A in transgenic peanut ( Arachishypogaea L) increases transpiration ef 1047297ciency under water-limiting conditionsPlant Cell Rep 2007262071ndash82
Bhatnagar-Mathur P Devi MJ Vadez V Sharma KK Differential antioxidative responses intransgenic peanut bear no relationship to their superior transpiration ef 1047297ciency underdrought stress J Plant Physiol 2009166(11)1207ndash17 doi101016jjplph200901001
Bidinger FR Serraj R Rizvi SMH Howarth C Yadav RS Hash CT Field evaluation of droughttolerance QTL effects on phenotype and adaptation in pearl millet [ Pennisetum glaucum(L) R Br] topcross hybrids Field Crops Res 200594(1)14ndash32
Bidinger FR Nepolean T Hash CT Yadav RS Howarth CJ Identi1047297cation of QTLs for grainyield of pearl millet (Pennisetum glaucum (L) R Br) in environments with variablemoisture during grain 1047297lling Crop Sci 200747969ndash80
Bowers JE Abbey C Anderson S Chang C Draye X A high-density geneticrecombination map of sequence-tagged sites for Sorghum as a framework forcomparative structural and evolutionary genomics of tropical grains and grassesGenetics 2003165367ndash86
Brick MA Ogg JB Singh SP Schwartz HF Johnson JJ Pastor-Corrales MA Registration of drought-tolerant rust-resistant high-yielding pinto bean germplasm lineCO46348 J Plant Registr 20082(2)120ndash4
Browne J Tunnacliffe A Burnell A Anhydrobiosis-plant desiccation gene found in anematode Nature 200241638
Busk PK Pages M Regulation of abscisic acid induced transcription Plant Mol Biol199837425ndash35
Carena MJ Wanner DW Development of genetically broad-based inbred lines of maizefor early-maturing (70-80RM) hybrids J Plant Registr 20093107ndash11
Cash SDBrucknerPL Wichman DMKephart KD Berg JEBoynerR et alRegistration of Willow Creek forage wheat J Plant Registr 20093(2)185ndash90
Cattivelli L Rizza F Badeck FW Mazzucotelli E Francia AMEM Mare AT et al Droughttolerance improvement in crop plants an integrated view from breeding to genomics
Field Crops Res 20081051-14Chen M Wang QY Cheng XG Xu ZS Li LC Ye XG et al GmDREB2 a soybean DRE-
binding transcription factor conferred drought and high-salt tolerance intransgenic plants Biochem Biophys Res Commun 2007353299ndash305
Cheng Z Targolli J Huang X Wu R Wheat LEA genes PMA80 and PMA1959 enhancedehydration tolerance of transgenic rice (Oryza sativa L) Mol Breed 20021071ndash82
ChoEK Hong ChB Over-expression of tobacco NtHSP70-1 contributes to drought-stresstolerance in plants Plant Cell Reports 200625349ndash58
Close TJ Dehydrins a commonality in the response of plants to dehydration and lowtemperature Physiol Plant 1997100291ndash6
Collins NC Tardieu F Tuberosa R Quantitative trait loci and crop performance underabiotic stress where do we stand Plant Physiol 2008147469ndash86
Concept Note Combining breeding and biotechnology to develop water ef 1047297cient maizefor Africa (WEMA) Afr Agric Technol Foundation
Courtois BShen L Petalcorin WCarandang S MauleonR LiZ Locating QTLs controllingconstitutive root traits in the rice population IAC 165-Co39 Euphytica 2003134335ndash45
Da-hong L Hui L Yang YL Ping-ping Z Jian-sheng L Down-regulated expression of RACK1geneby RNAinterference enhancesdroughttolerancein rice RiceSci 200916(1)14ndash20
Dalal M Tayal D Chinnusamy V Bansala KC Abiotic stress and ABA-inducible group 4 LEAfrom Brassicanapus plays a key rolein saltand droughttolerance J Biotechnol 2009139137ndash45
Ding Z Li S An X Liu X Qin H Wang D Transgenic expression of MYB15 confers enhancedsensitivity to abscisic acid and improved drought tolerance in Arabidopsis thaliana
J Genet Genomics 20093617ndash29Dubouzet JG Sakuma Y Ito Y Kasuga M Dubouzet EG Miura S et al OsDREB genes in
rice Oryza sativa L encode transcription activators that function in drought high-salt- and cold-responsive gene expression Plant J 200333751ndash63
El-Far IA Allan AY Responses of some wheat cultivars to sowing methods and droughtat different stages of growth Assuit J Agric Sci 199526(1)267ndash77
Eltayeb AE Kawano N Badawi GH Kaminaka H Sanekata T Shibahara T et alOverexpression of monodehydroascorbate reductase in transgenic tobacco confersenhanced tolerance to ozonesalt and polyethyleneglycol stresses Planta 2007225(5)1255ndash64
Falconer DS Introduction to quantitative genetics London New York Longman 1989Feng-ling FU Zhi-Lei F Shi-bing G Shu-feng Z Wan-chen L Evaluation and quantitative
inheritanceof several drought-relativetraits in maize Agric SciChina 20087(3)280ndash90Finkelstein R Gampala S Rock C Abscisic acid signaling in seeds and seedlings Plant
Cell 20021415ndash45Flowers TJ Improving crop salt tolerance J Exp Bot 200455307ndash19Fujita Y Fujita M Satoh R Maruyama K Parvez MM Seki M et al AREB1 is a transcription
Giraudat J Parcy F Bertauche N Gosti F Leung J Morris PC et al Current advances inabscisic acid action and signalling Plant Mol Biol 1994261557ndash77
Gorbalenya AE Koonin EV Helicases amino acid sequence comparisons and structurendashfunction relationships Curr Opin Struct Biol 19933419ndash29
Gosal SS Wani SH Kang MS Biotechnology and drought tolerance J Crop Improvement20092319ndash54
Gubis J Vaňkovaacute R Červenaacute V Draguacuteňovaacute M Hudcovicovaacute M Lichtnerovaacute H et alTransformedtobacco plants with increased toleranceto droughtSouthAfr J Bot200773505ndash11
GuoO Zhang J GaoQ Xing SLi F Wang WDrought tolerancethrough over-expressionof mono ubiquitin in transgenic tobacco J Plant Physiol 20081651745 ndash55
Haley SD Johnson JJ Peairs FB Quick JS Stromberger JA Clayshulte SR et al Registration of lsquoRipperrsquo wheat J Plant Registr 200711ndash6
Han SE Park SR Kwon HB Yi BY Lee GB Byun MO Genetic engineering of drought-resistant tobacco plants by introducingthe trehalose phosphorylase (TP) gene fromPleurotus sajor-caju Plant Cell Tissue Organ Cult 200582151ndash8
Harris K Klein R Mullet J Sorghum stay-green QTL individually reduces post-1047298oweringdrought-induced leaf senescence J Exp Bot 200758327ndash38
Hong BS Zong-Suo L Ming-An S LEA proteins in higher plants structure functiongene expression and regulation Colloids Surf B Biointerf 200545131ndash5
Howarth CJ Yadav RS Successful marker assisted selection for drought tolerance anddisease resistance in pearl milletIGER Innovations 2002
Humphreys MO Humphreys MW Breeding for stress resistance general principles InAshraf M Harris PJC editors Abiotic stresses plant resistance through breedingand molecular approaches 2005 p 19ndash46
Ingram J Bartels D The molecular basis of dehydration tolerance in plants Ann RevPlant Physiol Plant Mol Biol 199647377ndash403
Jaglo-Ottosen KR Gilmour SJ Zarka DG Schabenberger O Thomashow MF ArabidopsisCBF1 overexpression induces COR genes and enhances freezing tolerance Science1998280104ndash6
Jang IC Oh SJ Seo JS Choi WB Song SI Kim CH et al Expression of a bifunctional fusion of the Escherichia coli genes for trehalose-6-phosphate synthase and trehalose-6-phosphate phosphatase in transgenic rice plants increases trehalose accumulationandabiotic stresstolerancewithoutstuntinggrowth Plant Physiol2003131516ndash24
Jeanneau M Gerentes D Foueillassar X Zivy M Vidal J Toppan A et al Improvement of drought tolerance in maize towards the functional validation of the Zm-Asr1 gene andincrease of water use ef 1047297ciency by over-expressing C4-PEPC Biochimie 2002841127ndash35
Jenson KB Palazzo AJ Waldron BL Bushman BS Registration of lsquoFirstStrikersquo Slenderwheatgrass J Plant Registr 2007124ndash5
Joung-youn K Hyung-in C Min-young I Soo-young K Arabidopsis basic leucine zipperproteins that mediate stress-responsive abscisic acid signaling Plant Cell 20024343ndash57
Juskiw PE Helm JH Oro M Nyachiro JM Salmon DF Registration of lsquoBentleyrsquo barley
J Plant Registr 20093(2)119ndash23Kalamaki MS Alexandrou D Lazari D Merkouropoulos G Fotopoulos V Pateraki I et al
Over-expression of a tomato N-acetyl-L -glutamate synthase gene (SlNAGS1) in Arabidopsis thaliana results in high ornithine levels and increased tolerance in saltand drought stresses J Exp Bot 200960(6)1859ndash71
Kamoshita A Babu CR Boopathi NM Fukai S Phenotypic and genotypic analysis of drought-resistance traits for development of rice cultivars adapted to rainfedenvironments Field Crops Res 2008109(103)1-23 doi101016jfcr200806010
KarakasB Ozias-AkinsP Stushnoff C SuefferheldM Rieger M Salinityand drought toleranceof mannitol-accumulating transgenic tobacco Plant Cell Environ 199720609ndash16
Karim S Aronsson H Ericson H Pirhonen M Leyman B Welin B et al Improved droughttolerance without undesired side effects in transgenic plants producing trehalose PlantMol Biol 200764371ndash86
Kasuga M Liu Q Miura S Yamaguchi-Shinozaki K Shinozaki K Improving plant droughtsaltand freezing tolerance by gene transfer of a single stress-inducible transcription factorNat Biotechnol 199917287ndash91
Kindiger M Gaub H Hasegawac M Katsurab Y Ueyamad K Gotob S et al Registrationof lsquoNanryorsquo tall fescue Crop Sci 2006461815ndash6
Kong I Dong J HartGE Characteristics linkage mappositionsand allelic differentiationof Sorghum bicolour (L) Moench by DNA simple-sequence repeats (SSRs) TheorAppl Genet 2000101438ndash48
Kumar R Venuprasad R Atlin GN Genetic analysis of rainfed lowland rice droughttolerance under naturally-occurring stress in eastern India heritability and QTL effects Field Crops Res 200710342ndash52
La1047297tte HR Price AH Courtois B Yield response to water de1047297cit in an upland ricemapping population associations among traits and genetic markers Theor ApplGenet 20041091237ndash46
Lal S Gulyani V Khurana P Overexpressionof HVA1 gene frombarley generates tolerance tosalinity and water stress in transgenic mulberry (Morus indica) Transgenic Res 200817
(4)651ndash
63Lanceras J Pantuwan G Jongdee B Toojinda T Quantitative trait loci associated withdrought tolerance at reproductive stage in rice Plant Physiol 2004135384ndash99
Levi A Ovnat L Paterson AH Saranga Y Photosynthesis of cotton near-isogenic linesintrogressed with QTLs for productivity and drought related traits Plant Sci 2009a17788ndash96
Levi A Paterson AH Barak V Yakir D Wang B Chee PW et al Field evaluation of cottonnear-isogenic lines introgressed with QTLs for productivity and drought relatedtraits Mol Breed 2009b23179ndash95
Liu Q Kasuga M Sakuma Y Abe H Miura S Yamaguchi- Shinozaki K et al Twotranscription factors DREB1 and DREB2 with an EREBPAP2 DNA binding domainseparate two cellular signal transduction pathways in drought-and low-temper-ature-responsive gene expression respectively in Arabidopsis Plant Cell 1998101391ndash406
Liu X Hua X Guo J Qi D Wang L Liu Z et al Enhanced tolerance to drought stress intransgenic tobacco plants overexpressing Biotechnol Lett 2008301275ndash80
Liu X Wanga Z Wanga L Wua R Phillips J Deng X LEA 4 group genes from theresurrection plant Boea hygrometrica confer dehydration tolerance in transgenictobacco Plant Sci 200917690ndash8
Luchi S Kobayashi M Taji T Naramoto M Seki M Kato T et al Regulation of droughttolerance by gene manipulation of 9-cis-epoxycarotenoid dioxygenase a keyenzyme in abscisic acid biosynthesis in Arabidopsis Plant J 200127325ndash33
Luo Y Liu YB Dong YX Gao XQ Zhang YS Expression of a putative alfalfa helicase increasestolerance to abioticstress in Arabidopsis by enhancing the capacities for ROS scavengingand osmotic adjustment J Plant Physiol 2009166385ndash94
MacLean JLDaweDC Hardy B Hettel GPRiceAlmanac sourcebook for themostimportanteconomic activity on Earth3rd ed Wallingford England CABI Publishing 2002
Mansour MMF Nitrogen containing compounds and adaptation of plants to salinitystress Biol Plant 200043491ndash500
McKersie BD Bowley SR Harjanto E Leprince O Water-de1047297cit tolerance and 1047297eldperformance of transgenic alfalfa overexpressing superoxide dismutase Plant Physiol19961111177ndash81
McKersie BD Murnaghan J Bowley SR Manipulating freezing tolerance in transgenicplants Acta Physiol Plant 199719485ndash95
Miranda JA Avonce N Suaacuterez R Thevelein JM Dijck PV Iturriaga G A bifunctional TPS-TPP enzyme from yeast confers tolerance to multiple and extreme abiotic-stressconditions in transgenic Arabidopsis Planta 2007226(6)1411ndash21
Mohammadi M Taleei A Zeinali H Naghavi MR Ceccarelli S Grando Baum M QTL analysis for phenologic traits in doubled haploid population of barley Int J AgricBiol 20057(5)820ndash3
Mohanty A Kathuria H Ferjani A Sakamoto A Mohanty P Murata N et al Transgenicsof an elite indica rice variety Pusa Basmati 1 harbouring the codA gene are highlytolerant to salt stress Theor Appl Genet 200210651ndash7
Noaman MM El Sayad AA Asaad FA El Sherbini AM El Bawab AO El Moselhi MA et alRegistration of lsquoGiza 126rsquo barley Crop Sci 199535(6)1710
Noaman MM Ahmed IA El-Sayed AA Abo-El-Enin RA El-Gamal AS El-Sherbiny AM et alRegistration of lsquoGiza 2000rsquo drought-tolerant six-rowed barley for rainfed and newreclaimed areas in Egypt Crop Sci 200747440
Obert DE Evans CP Wesenberg DM Windes JM Erickson CA Jackson EW et alRegistration of lsquoLenetahrsquo spring barley J Plant Registr 20082(2)85ndash7
Oh SJ Jeong JS Kim EH Yi NR Yi SI Jang IC et al Matrix attachment region fromthe chicken lysozyme locus reduces variability in transgene expression andconfers copy number-dependence in transgenic rice plants Plant Cell Rep20054145ndash54
Oh SJ Kim YS Kwon C Park HK Jeong JS Kim JK Overexpression of the transcriptionfactor AP37 in rice improves grain yield under drought conditions Plant Physiol
20092191ndash
200 doi101104pp109137554Owttrim GW RNA helicases and abiotic stress Nucleic AcidsRes 200634(11)3220ndash30Park BJ Liu Z Kanno A Kameya T Increased tolerance to salt and water de1047297cit stress in
transgenic lettuce (Lactuca sativa L) by constitutive expression of LEA PlantGrowth Regul 2005a45165ndash71
Park BJ Liu Z Kanno A Kameya T Genetic improvement of Chinese cabbage for salt anddroughttolerance by constitutiveexpression of a B napus LEAgenePlantSci 2005b169553ndash8
Pastori GM Foyer CH Common componentsnetworks and pathways of crosstoleranceto stress The central role of ldquoredoxrdquo and abscisic acid-mediated controls PlantPhysiol 2002129460ndash8
PellegrineschiA ReynoldsM PachecoM Brito RMAlmeraya R Yamaguchi-ShinozakiK et alStress-induced expression in wheat of the Arabidopsis thaliana DREB1Agenedelayswaterstress symptoms under greenhouse conditions Genome 200447493ndash500
Perl A Perl-Treves R Galili S Aviv D Shalgi E Malkin S et al Enhanced oxidative-stressdefense in transgenic potato overexpressing tomato Cu Zn superoxide dismutaseTheor Appl Genet 199385568ndash76
Podlich DW Winkler CR Cooper M Mapping as you go an effective approach formarker-assisted selection of complex traits Crop Sci 2004441560ndash71
182 M Ashraf Biotechnology Advances 28 (2010) 169ndash183
Pratt RC Casey MA Registration of maize germplasm line Oh605 Crop Sci 2006461004ndash5Qin F Kakimoto M Sakuma Y Maruyama K Osakabe Y Tran LSP et al Regulation and
functional analysis of ZmDREB2A in response to drought and heat stress in Zea mays LPlant J 20075054ndash69
Quan R Shang M Zhang H Zhao Y Zhang J Engineering of enhanced glycinebetainesynthesis improves drought tolerance in maize Plant Biotechnol J 20042477ndash86
Quarrie SA Gulli M Calestani C Steed A Marmiroli N Location of a gene regulatingdrought-induced abscisic acid production on the long arm of chromosome 5A of wheat Theor Appl Genet 199489794ndash800
QuickJS Stromberger JA Clayshulte S Clifford B Johnson JJ Peairs FB et al Registrationof lsquoPrairie Redrsquo wheat Crop Sci 2001411362ndash3
RajaramS Prospects andpromise ofwheatbreedingin the21stcentury Euphytica 20011193-15Rajaram S Role of conventional plant breeding and biotechnology in future wheat
production Turk J Agric Forest 200529105ndash11Rami JF Dufour P Trouche G Fliedel G Mestres C Davrieux F et al Quantitative trait
loci for grain quality productivity morphological and agronomical traits insorghum (Sorghum bicolor L Moench) Theor Appl Genet 199897605ndash16
Rassaa N Salahb HBH Latiri K Thermal responses of durum wheat Triticum durum toearly water stress consequence on leaf and 1047298ower development Plant Biol Pathol2008331(5)363ndash71
Reddy LJ Nigam SN Rao RCN Reddy NS Registration of ICGV 87354 peanut germplasmwith drought tolerance and rust resistance Crop Sci 200141274ndash5
Ribaut JM Ragot M Marker-assisted selection to improve drought adaptation in maize thebackcross approach perspectives limitations and alternatives J Exp Bot 200658351ndash60
Robin S Pathan MS Courtois B La1047297tte R Carandang S Lanceras S et al Mappingosmotic adjustment in an advanced back-cross inbred population of rice Theor
Appl Genet 2003107(7)1288ndash96Romero C Belles JM Vaya JL Serrano R Culianez-Macia FA Expression of the yeast
trehalose-6-phosphate synthase gene in transgenic tobacco plants pleiotropicphenotypes include drought tolerance Planta 1997201293ndash7
Ronde JAD Cress WA Krugerd GHJ Strasserd RJ Van Staden J Photosynthetic responseof transgenic soybean plants containing an Arabidopsis P5CR gene during heat anddrought stress J Plant Physiol 20041611211ndash24
Salvi S Tuberosa R To clone or not to clone plant QTLs present and future challengesTrends Plant Sci 200510297ndash304
Sanchez AC Subudhi PK Rosenow DT Nguyen HT Mapping QTLs associated with droughtresistance in sorghum (Sorghum bicolor L Moench) Plant Mol Biol 200248713ndash26
Saranga Y Menz M Jiang CX Wright RJ Yakir D Paterson AH Genomic dissection of genotypetimesenvironment interactions conferring adaptation of cotton to aridconditions Genome Res 2001111988ndash95
Sari-Gorla M Krajewski P Di Fonzo N Villa M Frova C Genetic analysis of droughttolerance in maize by molecular markers II Plant height and 1047298owering Theor ApplGenet 199999289ndash95
Seki M Narusaka M Ishida J Nanjo T Fujita M Oono Y et al Monitoring the expressionpro1047297les of 7000 Arabidopsis genes under drought cold and high-salinity stressesusing a full-length cDNA microarray Plant J 200231279ndash92
Serraj R Sinclair TR Osmolyte accumulation can it really increase crop yield underdrought conditions Plant Cell Environ 200225333ndash41
Serraj R Krishnamurthy L Kashiwagi J Kumar J Chandra S Crouch JH Variation in roottraits of chickpea (Cicer arietinum L) grown under terminal drought Field CropsRes 200488115ndash27
Serraj R Hash CT Rizvi MHS Sharma A Yadav RS Bidinger FR Recent advances in marker-assisted selection for drought tolerance in pearl millet Plant Prod Sci 20058(3)334ndash7
Shen YG Du BX Zhang WK Zhang JS Chen SY AhCMO regulated by stresses in Atriplexhortensis can improve drought tolerance in transgenic tobacco Theor Appl Genet2002105815ndash21
Shinozaki K Yamaguchi-Shinozaki K Molecular responses to dehydration and lowtemperature differences and cross-talk between two stress signaling pathwaysCurr Opin Plant Biol 20003217ndash23
Singh KB Omar M Saxena MC Johansen C Registration of FLIP 87-59C a drought-tolerant chickpea germplasm line Crop Sci 199636(2)1ndash2
Singh SP Teran H Gutierrez JA Registration of SEA 5 and SEA 13 drought tolerant drybean germplasm Crop Sci 200141276ndash7
Singh SP Teran H Lema M Schwartz HF Miklas PN Registration of white moldresistant dry bean germplasm line A 195 J Plant Registr 2007162ndash3
Sivamani E Bahieldin A Wraith JM Improved biomass productivity and water useef 1047297ciency under water de1047297cit conditions in transgenic wheat constitutivelyexpressing the barley HVA1 gene Plant Sci 20001551ndash9
Skovmand B Reynolds MP DeLacy IH Searching genetic resources for physiologicaltraits with potential for increasing yield In Reynolds MP Ortiz-Monasterio IMcNab A editors Application of Physiology in Wheat Breeding 2001 p 17ndash28
Steele K Novel upland rice variety bred using marker-assisted selection and client-oriented breeding released in Jharkhand India Bangor University 2009
Steele KA Price AH Shashidar HE Witcombe JR Marker-assistedselection to introgressrice QTLs controlling root traitsinto an Indianupland rice variety Theor Appl Genet2006112208ndash21
Steele KA Virk DS Kumar R Prasad SC Witcombe JR Field evaluation of upland ricelines selected for QTLs controlling root traits Field Crops Res 2007101180 ndash6
Sunkar R Kapoor A Zhu JK Post transcriptional induction of two CuZn superoxidedismutase genes in Arabidopsis is mediated by down regulation of miR398 andimportant for oxidative stress tolerance Plant Cell 2006182051ndash65
Talame V Sanguineti MC Chiapparino E Bahri H Ben Salem M Forster BP et alIdenti1047297cation of Hordeum spontaneum QTL alleles improving 1047297eld performance of barley grown under rainfed conditions Ann Appl Bot 2004144309ndash20
Tanner NKCordinO BanroquesJ DoereM Linder PThe Q Motif a newlyidenti1047297ed motif in DEAD box helicases may regulate ATP binding and hydrolysis Mol Cell 200311127ndash38
Taramino G Tarchini R Ferrario S Lee M Pe ME Characterization and mapping of simplesequence repeats (SSRs) in Sorghum bicolor Theor Appl Genet 19979566ndash72
Teulat B Monneveux P Wery J Borriegraves C Souyris I Charrier A et al Relationshipsbetween relative water content andgrowth parameters in barleya QTLstudy NewPhytol 199713799-107
Thi Lang N Chi Buu B Fine mapping for drought tolerance in rice ( Oryza sativa L)Omonrice 2008169-15Till BJ Comai L Henikoff S Tillering and ecotillering for crop improvement In
Varshney RK Tuberosa R editors Genomics-assisted crop improvement genomicsapproaches and platforms vol 1 2007 p 333ndash50
Tondelli A Francia E Barabaschi D Aprile A Skinner JS Stockinger EJ et al Mappingregulatory genes as candidates for cold and drought stress tolerance in barleyTheor Appl Genet 2006112445ndash54
Tuberosa R Salvi S Genomics approaches to improve drought tolerance in cropsTrends Plant Sci 200611405ndash12
Valkoun JJ Wheat pre-breeding using wild progenitors Euphytica 200111917ndash23Vashisht AA Tuteja N Stress responsive DEAD-box helicases a new pathway to
engineer plant stress tolerance J Photochem Photobiol B Biol 200684150 ndash60Vendruscolo ECG Schuster I Pileggi M Scapim CA Molinari HBC Marur CJ et al Stress-
induced synthesis of proline confers tolerance to water de1047297cit in transgenic wheat J Plant Physiol 20071641367ndash76
Vienne D Leonardi A Damerval C Zivy M Genetics of proteome variation for QTL characterization application to drought stress responses in maize J Exp Bot 199950303ndash9
Villareal RL Mujeeb-Kazi A Rajaram S Toro ED Morphological variability in somesynthetic hexaploid wheats derived from Triticum turgidum times T tauschii J GenetBreed 1994487-16
Vinh NT Paterson AH Genome mapping and its implication for stress resistance inplants In Ashraf M Harris PJC editors Abiotic stresses plant resistance throughbreeding and molecular approaches 2005
Vinocur B Altman A Recent advances in engineering plant tolerance to abiotic stressachievements and limitations Curr Opin Biotechnol 200516123ndash32
Wang YJ Hao YJ Zhang ZG Chen T Zhang JS Chen SY Isolation of trehalose-6-phosphate phosphatase gene from tobacco and its functional analysis in yeast cells
J Plant Physiol 2005162215ndash23Wu R Garg A Engineering rice plants with trehalose-producing genes improves
tolerance to drought salt and low temperature ISB News Report 2003Xiao B Huang Y Tang N Xiong L Overexpression of LEA gene in rice improves drought
resistance under 1047297eld conditions Theor Appl Genet 200711535ndash46Xinglai P Sangang X Qiannying P Yinhong S Registration of lsquo Jinmai 50rsquo wheat Crop Sci
200646983ndash5Xiong L Lee H Ishitani M Zhu JK Regulation of osmotic stress-responsive gene
expression by the LOS6ABA1 locus in Arabidopsis J Biol Chem 20022778588ndash96Xu GW Magill CW Shertz KF Hart GE A RFLP linkage map of Sorghum bicolor (L)
Moench Theor Appl Genet 199489139ndash45Xu D Duan X Wang B Hong BHo THD Wu RExpression of a late embryogenesis abundant
protein gene HVA1 from barley confers tolerance to water de1047297cit and salt stress intransgenic rice Plant Physiol 1996110249ndash57
Yadav RS Hash CT Bidinger FR Devos KM Howarth CJ Genomic regions associatedwith grain yield and aspects of post-1047298owering drought tolerance in pearl milletacross stress environments and testers background Euphytica 2004136265ndash77
YamadaM MorishitaH Urano K Shiozaki N Yamaguchi-Shinozaki K ShinozakiK et alEffects of free proline accumulation in petunias under drought stress J Exp Bot2005561975ndash81
Yang WJ Rich PJ Axtell JD Wood KV Bonham CC Ejeta G et al Genotypic variation forglycinebetaine in sorghum Crop Sci 200343162ndash9
Yan-Ying QU Ping MU Xue-Qin L Yu-Xiu T Feng W Hong-Liang Z et al QTL mappingand correlations between leaf water potential and drought resistance in rice underupland and lowland environments Acta Agron Sin 200834(2)198ndash206
Ye XDWu XLZhaoH Frehner MNoumlsberger J Potrykus Iet al Altered fructan accumulationin transgenic Lolium multi 1047298orum plants expressing a Bacillus subtilis sacB gene Plant Cell
Rep 200120205ndash
12Zaharieva M Gaulin E Havaux M Acevedo E Monneveux P Drought and heat responses in
the wild wheat relative Aegilops geniculata Roth potential interest for wheatimprovement Crop Sci 2001411321ndash9
Zhang J Zheng HGAartiA Pantuwan G NguyenTT Tripathy JNet alLocatinggenomicregions associated with components of drought resistance in rice comparativemapping within and across species Theor Appl Genet 200110319ndash29
Zhang GH Su Q An LJ Wu S Characterization and expression of a vacuolar Na+H+
antiporter gene from the monocot halophyte Aeluropus littoralis Plant PhysiolBiochem 200846117ndash26
Zhao J QTLs for oil content and their relationships to other agronomic traits in anEuropeantimesChinese oilseed rape population Germany Diss Grorg-Agust Univer-sity of Goettingen 2002
Zhao J Ren W Zhi D Wang L Xia G Arabidopsis DREB1ACBF3 bestowed transgenic tallfescue increased tolerance to drought stress Plant Cell Rep 2007261521ndash8
Zhou W Li Y Zhao BC Ge RC Shen YZ Wang G Huang ZJ Over-expression of TaSTRG gene improves salt and drought tolerance in rice J Plant Physiol 200915(166)1660ndash70
and cost-effective Molecular mapping and analysis of QTL have been
carried out for a number of qualitative and quantitative traits including
stress tolerance which has undoubtedly resulted in a great magnitude
of knowledge and better understanding of the causal genetic phenom-
ena that regulate these traits However limited success has resulted by
using this knowledge to manipulate genes in an effective way for the
improvement of a speci1047297c trait in crops although some crop cultivars
developed through using these tools have performed extremely well
under1047297
eld stressconditions However there are a variety of reasons forthe limited success in terms of achieving enhanced drought tolerance
using the marker-assisted breeding approach For example a major
dif 1047297culty confronting the scientists is the challenge of precise QTL
identi1047297cation In addition a substantial genetictimes environment interac-
tion inconsistent repeatability large number of genes regulating yield
and invalid use of mapping populations have hampered the pursuits
involving mapping of QTL for enhanced drought stress tolerance
Despite these other factors also impede the application of QTL for
genetic improvement of a trait For example due to unfavorable
epistatic interaction it is hard to transfer the effects of a desired allele to
an elite background material (Podlich et al 2004 Collins et al 2008)
Furthermore in some cases QTL from a speci1047297c background do not
show signi1047297cant effects or cease completely in different backgrounds
even under analogous growth conditions (Cho and Hong 2006 Collins
et al 2008) This makes the utilization of QTL more intricate
Transformation of the knowledge acquired from QTL-oriented molec-
ular studies into a well-de1047297ned upshot for the stakeholders is one of the
key challenges confronting the breeders Certainly a multidisciplinary
approach including more speci1047297cally the identi1047297cation of QTL mediated
signal transduction in response to stresses needs to be adopted
Furthermore although QTL cloning procedures are unwieldy they can
lead to a meaningful outcome because a cloned QTL can offer a reliable
markerfor MABas well as furnish an outlinefor thedetection of superior
allelic variants in crop species (Till et al 2007)
Genetic engineering (Transgenic approach) offers a promise whereby
one can expect a substantial improvement in a desired trait within the
shortest stretch of time Engineering genes encoding organic osmolytes
plant growth regulators antioxidants late embryogenesis abundant pro-
teins and transcription factors has resulted into transgenic lines whichhave out-performed under controlled stress conditions In most cases the
claims made by the researchers in terms of performance of transgenic
lines tested under controlled conditions are considerably in1047298ated
However with the exception of a very few notable cases most of the
transgenic lines of differentcrops have been rarely1047297eld-tested Thushow
far these transgenic lines perform under 1047297eld stress conditions is not
known because under natural1047297eld conditions a linecultivar has to face a
multitude of environmental factors other than the drought stress Thus
appraisal of performance of a transgenic line under natural 1047297eld con-
ditionsis mandatory forthe stakeholdersperspective It is also imperative
to note that most of the drought tolerant transgenic lines have been
developed using a single gene transformation which may not be as
productive as if it had been developed using transformation of many
genes Thus transferring a number of prominent genes effectively in-volved in stress tolerance to transgenic plants seems to be a logical
approach Although a large number of genes appear to be involved in
stress tolerance and most of them have been fully characterized the
function of many of them in the mechanism of stress tolerance is yet to be
investigated
References
Abebe T Guenzi AC Martin B Cushman JC Tolerance of mannitol-accumulatingtransgenic wheat to water stress and salinity Plant Physiol 20031311748ndash55
Ashraf M Breeding for salinity tolerance in plants Crit Rev Plant Sci 19941317 ndash42Ashraf M Akram NA Improving salinity tolerance of plants through conventional
breeding and genetic engineering an analytical comparison Biotechnol Adv200927744ndash52 doi101016jbiotechadv200905026
Ashraf M Foolad MR Roles of glycinebetaine and proline in improving plant abioticstress resistance Environ Exp Bot 200759206ndash16
Ashraf M Athar HR Harris PJC Kwon TR Some prospective strategies for improvingcrop salt tolerance Adv Agron 20089745-110
Asins MJ Present and future of quantitative trait locus analysis in plant breeding PlantBreed 2002121281ndash91
Babu RC Nguyen BD Chamarerk V Shanmugasundaram P Chezhian P Jeyaprakash P et alGeneticanalysis of droughtresistance in riceby molecular markers association betweensecondary traits and 1047297eld performance Crop Sci 2003431457ndash69
Babu RC Zhang J Blum A Ho THD Wu R Nguyen HT HVA1 a LEA gene from barleyconfers dehydration tolerance in transgenic rice( Oryzasativa L)via cell membraneprotection Plant Sci 2004166855ndash62
Badawi GH Kawano N Yamauchi Y Over-expression of ascorbate peroxidase in
tobacco chloroplasts enhances the tolerance to salt stress and water de1047297cit PhysiolPlant 2004121231ndash8Badu-Apraku B Yallou CG Registration of striga-resistant and drought tolerant tropical
early maize populations TZE-W Pop DT STR C4 and TZE-Y Pop DT STR C4 J PlantRegistr 20093(1)86ndash90
Badu-Apraku B Menkir A Kling JG Fakorede MAB Registration of 16 striga resistantearly maturing tropical maize inbred lines Crop Sci 2006461410ndash1
Baenziger PS Beecher B Graybosch RA Ibrahim AMH Baltensperger DD Nelson LA et alRegistration of lsquoNEO1643rsquo wheat J Plant Registr 20082(1)36ndash42
Baumlnziger M Setimela PS Hodson D Vivek B Breeding for improved drought tolerance inmaize adapted to southern Africa Proceedings of the 4th International Crop ScienceCongress Brisbane Australia Published on CDROM 2004 26 Sep ndash 1 Oct
Bartels D Sunkar R Drought and salt tolerance in plants Crit Rev Plant Sci 20052423 ndash58Baum M Grandol S Backes G Jahoor A Sabbagh A Ceccarelli S QTLs for agronomic
traits in the Mediterranean environment identi1047297ed in recombinant inbred lines of the cross lsquoArtarsquo H spontaneum 41-1 Theor Appl Genet 20031071215ndash25
Bergman JW Riveland NR Flynn CR Carlson GR Wichman DM Registration of lsquoMorlinrsquo
saf 1047298ower Crop Sci 2001411640Bernier J Kumar A Venuprasad R Spaner D Atlin G A large-effect QTL for
grain yield under reproductive-stage drought stress in upland rice Crop Sci200747507ndash18
BernierJ Kumar A SerrajR SpanerD Atlin G Review breeding uplandrice fordroughtresistance J Sci Food Agric 200888927ndash39
Bernier J Serraj R Kumar A Venuprasad R Impa S Gowdaa RPV et al The large-effectdrought-resistance QTL qtl121 increases water uptake in upland rice Field CropsRes 200911039ndash46
Bhatnagar-Mathur P ReddyDS Lavanya M Yamaguchi-Shinozaki K Sharma KK Stress-inducible expression of Arabidopsis thaliana DREB1A in transgenic peanut ( Arachishypogaea L) increases transpiration ef 1047297ciency under water-limiting conditionsPlant Cell Rep 2007262071ndash82
Bhatnagar-Mathur P Devi MJ Vadez V Sharma KK Differential antioxidative responses intransgenic peanut bear no relationship to their superior transpiration ef 1047297ciency underdrought stress J Plant Physiol 2009166(11)1207ndash17 doi101016jjplph200901001
Bidinger FR Serraj R Rizvi SMH Howarth C Yadav RS Hash CT Field evaluation of droughttolerance QTL effects on phenotype and adaptation in pearl millet [ Pennisetum glaucum(L) R Br] topcross hybrids Field Crops Res 200594(1)14ndash32
Bidinger FR Nepolean T Hash CT Yadav RS Howarth CJ Identi1047297cation of QTLs for grainyield of pearl millet (Pennisetum glaucum (L) R Br) in environments with variablemoisture during grain 1047297lling Crop Sci 200747969ndash80
Bowers JE Abbey C Anderson S Chang C Draye X A high-density geneticrecombination map of sequence-tagged sites for Sorghum as a framework forcomparative structural and evolutionary genomics of tropical grains and grassesGenetics 2003165367ndash86
Brick MA Ogg JB Singh SP Schwartz HF Johnson JJ Pastor-Corrales MA Registration of drought-tolerant rust-resistant high-yielding pinto bean germplasm lineCO46348 J Plant Registr 20082(2)120ndash4
Browne J Tunnacliffe A Burnell A Anhydrobiosis-plant desiccation gene found in anematode Nature 200241638
Busk PK Pages M Regulation of abscisic acid induced transcription Plant Mol Biol199837425ndash35
Carena MJ Wanner DW Development of genetically broad-based inbred lines of maizefor early-maturing (70-80RM) hybrids J Plant Registr 20093107ndash11
Cash SDBrucknerPL Wichman DMKephart KD Berg JEBoynerR et alRegistration of Willow Creek forage wheat J Plant Registr 20093(2)185ndash90
Cattivelli L Rizza F Badeck FW Mazzucotelli E Francia AMEM Mare AT et al Droughttolerance improvement in crop plants an integrated view from breeding to genomics
Field Crops Res 20081051-14Chen M Wang QY Cheng XG Xu ZS Li LC Ye XG et al GmDREB2 a soybean DRE-
binding transcription factor conferred drought and high-salt tolerance intransgenic plants Biochem Biophys Res Commun 2007353299ndash305
Cheng Z Targolli J Huang X Wu R Wheat LEA genes PMA80 and PMA1959 enhancedehydration tolerance of transgenic rice (Oryza sativa L) Mol Breed 20021071ndash82
ChoEK Hong ChB Over-expression of tobacco NtHSP70-1 contributes to drought-stresstolerance in plants Plant Cell Reports 200625349ndash58
Close TJ Dehydrins a commonality in the response of plants to dehydration and lowtemperature Physiol Plant 1997100291ndash6
Collins NC Tardieu F Tuberosa R Quantitative trait loci and crop performance underabiotic stress where do we stand Plant Physiol 2008147469ndash86
Concept Note Combining breeding and biotechnology to develop water ef 1047297cient maizefor Africa (WEMA) Afr Agric Technol Foundation
Courtois BShen L Petalcorin WCarandang S MauleonR LiZ Locating QTLs controllingconstitutive root traits in the rice population IAC 165-Co39 Euphytica 2003134335ndash45
Da-hong L Hui L Yang YL Ping-ping Z Jian-sheng L Down-regulated expression of RACK1geneby RNAinterference enhancesdroughttolerancein rice RiceSci 200916(1)14ndash20
Dalal M Tayal D Chinnusamy V Bansala KC Abiotic stress and ABA-inducible group 4 LEAfrom Brassicanapus plays a key rolein saltand droughttolerance J Biotechnol 2009139137ndash45
Ding Z Li S An X Liu X Qin H Wang D Transgenic expression of MYB15 confers enhancedsensitivity to abscisic acid and improved drought tolerance in Arabidopsis thaliana
J Genet Genomics 20093617ndash29Dubouzet JG Sakuma Y Ito Y Kasuga M Dubouzet EG Miura S et al OsDREB genes in
rice Oryza sativa L encode transcription activators that function in drought high-salt- and cold-responsive gene expression Plant J 200333751ndash63
El-Far IA Allan AY Responses of some wheat cultivars to sowing methods and droughtat different stages of growth Assuit J Agric Sci 199526(1)267ndash77
Eltayeb AE Kawano N Badawi GH Kaminaka H Sanekata T Shibahara T et alOverexpression of monodehydroascorbate reductase in transgenic tobacco confersenhanced tolerance to ozonesalt and polyethyleneglycol stresses Planta 2007225(5)1255ndash64
Falconer DS Introduction to quantitative genetics London New York Longman 1989Feng-ling FU Zhi-Lei F Shi-bing G Shu-feng Z Wan-chen L Evaluation and quantitative
inheritanceof several drought-relativetraits in maize Agric SciChina 20087(3)280ndash90Finkelstein R Gampala S Rock C Abscisic acid signaling in seeds and seedlings Plant
Cell 20021415ndash45Flowers TJ Improving crop salt tolerance J Exp Bot 200455307ndash19Fujita Y Fujita M Satoh R Maruyama K Parvez MM Seki M et al AREB1 is a transcription
Giraudat J Parcy F Bertauche N Gosti F Leung J Morris PC et al Current advances inabscisic acid action and signalling Plant Mol Biol 1994261557ndash77
Gorbalenya AE Koonin EV Helicases amino acid sequence comparisons and structurendashfunction relationships Curr Opin Struct Biol 19933419ndash29
Gosal SS Wani SH Kang MS Biotechnology and drought tolerance J Crop Improvement20092319ndash54
Gubis J Vaňkovaacute R Červenaacute V Draguacuteňovaacute M Hudcovicovaacute M Lichtnerovaacute H et alTransformedtobacco plants with increased toleranceto droughtSouthAfr J Bot200773505ndash11
GuoO Zhang J GaoQ Xing SLi F Wang WDrought tolerancethrough over-expressionof mono ubiquitin in transgenic tobacco J Plant Physiol 20081651745 ndash55
Haley SD Johnson JJ Peairs FB Quick JS Stromberger JA Clayshulte SR et al Registration of lsquoRipperrsquo wheat J Plant Registr 200711ndash6
Han SE Park SR Kwon HB Yi BY Lee GB Byun MO Genetic engineering of drought-resistant tobacco plants by introducingthe trehalose phosphorylase (TP) gene fromPleurotus sajor-caju Plant Cell Tissue Organ Cult 200582151ndash8
Harris K Klein R Mullet J Sorghum stay-green QTL individually reduces post-1047298oweringdrought-induced leaf senescence J Exp Bot 200758327ndash38
Hong BS Zong-Suo L Ming-An S LEA proteins in higher plants structure functiongene expression and regulation Colloids Surf B Biointerf 200545131ndash5
Howarth CJ Yadav RS Successful marker assisted selection for drought tolerance anddisease resistance in pearl milletIGER Innovations 2002
Humphreys MO Humphreys MW Breeding for stress resistance general principles InAshraf M Harris PJC editors Abiotic stresses plant resistance through breedingand molecular approaches 2005 p 19ndash46
Ingram J Bartels D The molecular basis of dehydration tolerance in plants Ann RevPlant Physiol Plant Mol Biol 199647377ndash403
Jaglo-Ottosen KR Gilmour SJ Zarka DG Schabenberger O Thomashow MF ArabidopsisCBF1 overexpression induces COR genes and enhances freezing tolerance Science1998280104ndash6
Jang IC Oh SJ Seo JS Choi WB Song SI Kim CH et al Expression of a bifunctional fusion of the Escherichia coli genes for trehalose-6-phosphate synthase and trehalose-6-phosphate phosphatase in transgenic rice plants increases trehalose accumulationandabiotic stresstolerancewithoutstuntinggrowth Plant Physiol2003131516ndash24
Jeanneau M Gerentes D Foueillassar X Zivy M Vidal J Toppan A et al Improvement of drought tolerance in maize towards the functional validation of the Zm-Asr1 gene andincrease of water use ef 1047297ciency by over-expressing C4-PEPC Biochimie 2002841127ndash35
Jenson KB Palazzo AJ Waldron BL Bushman BS Registration of lsquoFirstStrikersquo Slenderwheatgrass J Plant Registr 2007124ndash5
Joung-youn K Hyung-in C Min-young I Soo-young K Arabidopsis basic leucine zipperproteins that mediate stress-responsive abscisic acid signaling Plant Cell 20024343ndash57
Juskiw PE Helm JH Oro M Nyachiro JM Salmon DF Registration of lsquoBentleyrsquo barley
J Plant Registr 20093(2)119ndash23Kalamaki MS Alexandrou D Lazari D Merkouropoulos G Fotopoulos V Pateraki I et al
Over-expression of a tomato N-acetyl-L -glutamate synthase gene (SlNAGS1) in Arabidopsis thaliana results in high ornithine levels and increased tolerance in saltand drought stresses J Exp Bot 200960(6)1859ndash71
Kamoshita A Babu CR Boopathi NM Fukai S Phenotypic and genotypic analysis of drought-resistance traits for development of rice cultivars adapted to rainfedenvironments Field Crops Res 2008109(103)1-23 doi101016jfcr200806010
KarakasB Ozias-AkinsP Stushnoff C SuefferheldM Rieger M Salinityand drought toleranceof mannitol-accumulating transgenic tobacco Plant Cell Environ 199720609ndash16
Karim S Aronsson H Ericson H Pirhonen M Leyman B Welin B et al Improved droughttolerance without undesired side effects in transgenic plants producing trehalose PlantMol Biol 200764371ndash86
Kasuga M Liu Q Miura S Yamaguchi-Shinozaki K Shinozaki K Improving plant droughtsaltand freezing tolerance by gene transfer of a single stress-inducible transcription factorNat Biotechnol 199917287ndash91
Kindiger M Gaub H Hasegawac M Katsurab Y Ueyamad K Gotob S et al Registrationof lsquoNanryorsquo tall fescue Crop Sci 2006461815ndash6
Kong I Dong J HartGE Characteristics linkage mappositionsand allelic differentiationof Sorghum bicolour (L) Moench by DNA simple-sequence repeats (SSRs) TheorAppl Genet 2000101438ndash48
Kumar R Venuprasad R Atlin GN Genetic analysis of rainfed lowland rice droughttolerance under naturally-occurring stress in eastern India heritability and QTL effects Field Crops Res 200710342ndash52
La1047297tte HR Price AH Courtois B Yield response to water de1047297cit in an upland ricemapping population associations among traits and genetic markers Theor ApplGenet 20041091237ndash46
Lal S Gulyani V Khurana P Overexpressionof HVA1 gene frombarley generates tolerance tosalinity and water stress in transgenic mulberry (Morus indica) Transgenic Res 200817
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63Lanceras J Pantuwan G Jongdee B Toojinda T Quantitative trait loci associated withdrought tolerance at reproductive stage in rice Plant Physiol 2004135384ndash99
Levi A Ovnat L Paterson AH Saranga Y Photosynthesis of cotton near-isogenic linesintrogressed with QTLs for productivity and drought related traits Plant Sci 2009a17788ndash96
Levi A Paterson AH Barak V Yakir D Wang B Chee PW et al Field evaluation of cottonnear-isogenic lines introgressed with QTLs for productivity and drought relatedtraits Mol Breed 2009b23179ndash95
Liu Q Kasuga M Sakuma Y Abe H Miura S Yamaguchi- Shinozaki K et al Twotranscription factors DREB1 and DREB2 with an EREBPAP2 DNA binding domainseparate two cellular signal transduction pathways in drought-and low-temper-ature-responsive gene expression respectively in Arabidopsis Plant Cell 1998101391ndash406
Liu X Hua X Guo J Qi D Wang L Liu Z et al Enhanced tolerance to drought stress intransgenic tobacco plants overexpressing Biotechnol Lett 2008301275ndash80
Liu X Wanga Z Wanga L Wua R Phillips J Deng X LEA 4 group genes from theresurrection plant Boea hygrometrica confer dehydration tolerance in transgenictobacco Plant Sci 200917690ndash8
Luchi S Kobayashi M Taji T Naramoto M Seki M Kato T et al Regulation of droughttolerance by gene manipulation of 9-cis-epoxycarotenoid dioxygenase a keyenzyme in abscisic acid biosynthesis in Arabidopsis Plant J 200127325ndash33
Luo Y Liu YB Dong YX Gao XQ Zhang YS Expression of a putative alfalfa helicase increasestolerance to abioticstress in Arabidopsis by enhancing the capacities for ROS scavengingand osmotic adjustment J Plant Physiol 2009166385ndash94
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Mansour MMF Nitrogen containing compounds and adaptation of plants to salinitystress Biol Plant 200043491ndash500
McKersie BD Bowley SR Harjanto E Leprince O Water-de1047297cit tolerance and 1047297eldperformance of transgenic alfalfa overexpressing superoxide dismutase Plant Physiol19961111177ndash81
McKersie BD Murnaghan J Bowley SR Manipulating freezing tolerance in transgenicplants Acta Physiol Plant 199719485ndash95
Miranda JA Avonce N Suaacuterez R Thevelein JM Dijck PV Iturriaga G A bifunctional TPS-TPP enzyme from yeast confers tolerance to multiple and extreme abiotic-stressconditions in transgenic Arabidopsis Planta 2007226(6)1411ndash21
Mohammadi M Taleei A Zeinali H Naghavi MR Ceccarelli S Grando Baum M QTL analysis for phenologic traits in doubled haploid population of barley Int J AgricBiol 20057(5)820ndash3
Mohanty A Kathuria H Ferjani A Sakamoto A Mohanty P Murata N et al Transgenicsof an elite indica rice variety Pusa Basmati 1 harbouring the codA gene are highlytolerant to salt stress Theor Appl Genet 200210651ndash7
Noaman MM El Sayad AA Asaad FA El Sherbini AM El Bawab AO El Moselhi MA et alRegistration of lsquoGiza 126rsquo barley Crop Sci 199535(6)1710
Noaman MM Ahmed IA El-Sayed AA Abo-El-Enin RA El-Gamal AS El-Sherbiny AM et alRegistration of lsquoGiza 2000rsquo drought-tolerant six-rowed barley for rainfed and newreclaimed areas in Egypt Crop Sci 200747440
Obert DE Evans CP Wesenberg DM Windes JM Erickson CA Jackson EW et alRegistration of lsquoLenetahrsquo spring barley J Plant Registr 20082(2)85ndash7
Oh SJ Jeong JS Kim EH Yi NR Yi SI Jang IC et al Matrix attachment region fromthe chicken lysozyme locus reduces variability in transgene expression andconfers copy number-dependence in transgenic rice plants Plant Cell Rep20054145ndash54
Oh SJ Kim YS Kwon C Park HK Jeong JS Kim JK Overexpression of the transcriptionfactor AP37 in rice improves grain yield under drought conditions Plant Physiol
20092191ndash
200 doi101104pp109137554Owttrim GW RNA helicases and abiotic stress Nucleic AcidsRes 200634(11)3220ndash30Park BJ Liu Z Kanno A Kameya T Increased tolerance to salt and water de1047297cit stress in
transgenic lettuce (Lactuca sativa L) by constitutive expression of LEA PlantGrowth Regul 2005a45165ndash71
Park BJ Liu Z Kanno A Kameya T Genetic improvement of Chinese cabbage for salt anddroughttolerance by constitutiveexpression of a B napus LEAgenePlantSci 2005b169553ndash8
Pastori GM Foyer CH Common componentsnetworks and pathways of crosstoleranceto stress The central role of ldquoredoxrdquo and abscisic acid-mediated controls PlantPhysiol 2002129460ndash8
PellegrineschiA ReynoldsM PachecoM Brito RMAlmeraya R Yamaguchi-ShinozakiK et alStress-induced expression in wheat of the Arabidopsis thaliana DREB1Agenedelayswaterstress symptoms under greenhouse conditions Genome 200447493ndash500
Perl A Perl-Treves R Galili S Aviv D Shalgi E Malkin S et al Enhanced oxidative-stressdefense in transgenic potato overexpressing tomato Cu Zn superoxide dismutaseTheor Appl Genet 199385568ndash76
Podlich DW Winkler CR Cooper M Mapping as you go an effective approach formarker-assisted selection of complex traits Crop Sci 2004441560ndash71
182 M Ashraf Biotechnology Advances 28 (2010) 169ndash183
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functional analysis of ZmDREB2A in response to drought and heat stress in Zea mays LPlant J 20075054ndash69
Quan R Shang M Zhang H Zhao Y Zhang J Engineering of enhanced glycinebetainesynthesis improves drought tolerance in maize Plant Biotechnol J 20042477ndash86
Quarrie SA Gulli M Calestani C Steed A Marmiroli N Location of a gene regulatingdrought-induced abscisic acid production on the long arm of chromosome 5A of wheat Theor Appl Genet 199489794ndash800
QuickJS Stromberger JA Clayshulte S Clifford B Johnson JJ Peairs FB et al Registrationof lsquoPrairie Redrsquo wheat Crop Sci 2001411362ndash3
RajaramS Prospects andpromise ofwheatbreedingin the21stcentury Euphytica 20011193-15Rajaram S Role of conventional plant breeding and biotechnology in future wheat
production Turk J Agric Forest 200529105ndash11Rami JF Dufour P Trouche G Fliedel G Mestres C Davrieux F et al Quantitative trait
loci for grain quality productivity morphological and agronomical traits insorghum (Sorghum bicolor L Moench) Theor Appl Genet 199897605ndash16
Rassaa N Salahb HBH Latiri K Thermal responses of durum wheat Triticum durum toearly water stress consequence on leaf and 1047298ower development Plant Biol Pathol2008331(5)363ndash71
Reddy LJ Nigam SN Rao RCN Reddy NS Registration of ICGV 87354 peanut germplasmwith drought tolerance and rust resistance Crop Sci 200141274ndash5
Ribaut JM Ragot M Marker-assisted selection to improve drought adaptation in maize thebackcross approach perspectives limitations and alternatives J Exp Bot 200658351ndash60
Robin S Pathan MS Courtois B La1047297tte R Carandang S Lanceras S et al Mappingosmotic adjustment in an advanced back-cross inbred population of rice Theor
Appl Genet 2003107(7)1288ndash96Romero C Belles JM Vaya JL Serrano R Culianez-Macia FA Expression of the yeast
trehalose-6-phosphate synthase gene in transgenic tobacco plants pleiotropicphenotypes include drought tolerance Planta 1997201293ndash7
Ronde JAD Cress WA Krugerd GHJ Strasserd RJ Van Staden J Photosynthetic responseof transgenic soybean plants containing an Arabidopsis P5CR gene during heat anddrought stress J Plant Physiol 20041611211ndash24
Salvi S Tuberosa R To clone or not to clone plant QTLs present and future challengesTrends Plant Sci 200510297ndash304
Sanchez AC Subudhi PK Rosenow DT Nguyen HT Mapping QTLs associated with droughtresistance in sorghum (Sorghum bicolor L Moench) Plant Mol Biol 200248713ndash26
Saranga Y Menz M Jiang CX Wright RJ Yakir D Paterson AH Genomic dissection of genotypetimesenvironment interactions conferring adaptation of cotton to aridconditions Genome Res 2001111988ndash95
Sari-Gorla M Krajewski P Di Fonzo N Villa M Frova C Genetic analysis of droughttolerance in maize by molecular markers II Plant height and 1047298owering Theor ApplGenet 199999289ndash95
Seki M Narusaka M Ishida J Nanjo T Fujita M Oono Y et al Monitoring the expressionpro1047297les of 7000 Arabidopsis genes under drought cold and high-salinity stressesusing a full-length cDNA microarray Plant J 200231279ndash92
Serraj R Sinclair TR Osmolyte accumulation can it really increase crop yield underdrought conditions Plant Cell Environ 200225333ndash41
Serraj R Krishnamurthy L Kashiwagi J Kumar J Chandra S Crouch JH Variation in roottraits of chickpea (Cicer arietinum L) grown under terminal drought Field CropsRes 200488115ndash27
Serraj R Hash CT Rizvi MHS Sharma A Yadav RS Bidinger FR Recent advances in marker-assisted selection for drought tolerance in pearl millet Plant Prod Sci 20058(3)334ndash7
Shen YG Du BX Zhang WK Zhang JS Chen SY AhCMO regulated by stresses in Atriplexhortensis can improve drought tolerance in transgenic tobacco Theor Appl Genet2002105815ndash21
Shinozaki K Yamaguchi-Shinozaki K Molecular responses to dehydration and lowtemperature differences and cross-talk between two stress signaling pathwaysCurr Opin Plant Biol 20003217ndash23
Singh KB Omar M Saxena MC Johansen C Registration of FLIP 87-59C a drought-tolerant chickpea germplasm line Crop Sci 199636(2)1ndash2
Singh SP Teran H Gutierrez JA Registration of SEA 5 and SEA 13 drought tolerant drybean germplasm Crop Sci 200141276ndash7
Singh SP Teran H Lema M Schwartz HF Miklas PN Registration of white moldresistant dry bean germplasm line A 195 J Plant Registr 2007162ndash3
Sivamani E Bahieldin A Wraith JM Improved biomass productivity and water useef 1047297ciency under water de1047297cit conditions in transgenic wheat constitutivelyexpressing the barley HVA1 gene Plant Sci 20001551ndash9
Skovmand B Reynolds MP DeLacy IH Searching genetic resources for physiologicaltraits with potential for increasing yield In Reynolds MP Ortiz-Monasterio IMcNab A editors Application of Physiology in Wheat Breeding 2001 p 17ndash28
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Steele KA Price AH Shashidar HE Witcombe JR Marker-assistedselection to introgressrice QTLs controlling root traitsinto an Indianupland rice variety Theor Appl Genet2006112208ndash21
Steele KA Virk DS Kumar R Prasad SC Witcombe JR Field evaluation of upland ricelines selected for QTLs controlling root traits Field Crops Res 2007101180 ndash6
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Talame V Sanguineti MC Chiapparino E Bahri H Ben Salem M Forster BP et alIdenti1047297cation of Hordeum spontaneum QTL alleles improving 1047297eld performance of barley grown under rainfed conditions Ann Appl Bot 2004144309ndash20
Tanner NKCordinO BanroquesJ DoereM Linder PThe Q Motif a newlyidenti1047297ed motif in DEAD box helicases may regulate ATP binding and hydrolysis Mol Cell 200311127ndash38
Taramino G Tarchini R Ferrario S Lee M Pe ME Characterization and mapping of simplesequence repeats (SSRs) in Sorghum bicolor Theor Appl Genet 19979566ndash72
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Thi Lang N Chi Buu B Fine mapping for drought tolerance in rice ( Oryza sativa L)Omonrice 2008169-15Till BJ Comai L Henikoff S Tillering and ecotillering for crop improvement In
Varshney RK Tuberosa R editors Genomics-assisted crop improvement genomicsapproaches and platforms vol 1 2007 p 333ndash50
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Tuberosa R Salvi S Genomics approaches to improve drought tolerance in cropsTrends Plant Sci 200611405ndash12
Valkoun JJ Wheat pre-breeding using wild progenitors Euphytica 200111917ndash23Vashisht AA Tuteja N Stress responsive DEAD-box helicases a new pathway to
engineer plant stress tolerance J Photochem Photobiol B Biol 200684150 ndash60Vendruscolo ECG Schuster I Pileggi M Scapim CA Molinari HBC Marur CJ et al Stress-
induced synthesis of proline confers tolerance to water de1047297cit in transgenic wheat J Plant Physiol 20071641367ndash76
Vienne D Leonardi A Damerval C Zivy M Genetics of proteome variation for QTL characterization application to drought stress responses in maize J Exp Bot 199950303ndash9
Villareal RL Mujeeb-Kazi A Rajaram S Toro ED Morphological variability in somesynthetic hexaploid wheats derived from Triticum turgidum times T tauschii J GenetBreed 1994487-16
Vinh NT Paterson AH Genome mapping and its implication for stress resistance inplants In Ashraf M Harris PJC editors Abiotic stresses plant resistance throughbreeding and molecular approaches 2005
Vinocur B Altman A Recent advances in engineering plant tolerance to abiotic stressachievements and limitations Curr Opin Biotechnol 200516123ndash32
Wang YJ Hao YJ Zhang ZG Chen T Zhang JS Chen SY Isolation of trehalose-6-phosphate phosphatase gene from tobacco and its functional analysis in yeast cells
J Plant Physiol 2005162215ndash23Wu R Garg A Engineering rice plants with trehalose-producing genes improves
tolerance to drought salt and low temperature ISB News Report 2003Xiao B Huang Y Tang N Xiong L Overexpression of LEA gene in rice improves drought
resistance under 1047297eld conditions Theor Appl Genet 200711535ndash46Xinglai P Sangang X Qiannying P Yinhong S Registration of lsquo Jinmai 50rsquo wheat Crop Sci
200646983ndash5Xiong L Lee H Ishitani M Zhu JK Regulation of osmotic stress-responsive gene
expression by the LOS6ABA1 locus in Arabidopsis J Biol Chem 20022778588ndash96Xu GW Magill CW Shertz KF Hart GE A RFLP linkage map of Sorghum bicolor (L)
Moench Theor Appl Genet 199489139ndash45Xu D Duan X Wang B Hong BHo THD Wu RExpression of a late embryogenesis abundant
protein gene HVA1 from barley confers tolerance to water de1047297cit and salt stress intransgenic rice Plant Physiol 1996110249ndash57
Yadav RS Hash CT Bidinger FR Devos KM Howarth CJ Genomic regions associatedwith grain yield and aspects of post-1047298owering drought tolerance in pearl milletacross stress environments and testers background Euphytica 2004136265ndash77
YamadaM MorishitaH Urano K Shiozaki N Yamaguchi-Shinozaki K ShinozakiK et alEffects of free proline accumulation in petunias under drought stress J Exp Bot2005561975ndash81
Yang WJ Rich PJ Axtell JD Wood KV Bonham CC Ejeta G et al Genotypic variation forglycinebetaine in sorghum Crop Sci 200343162ndash9
Yan-Ying QU Ping MU Xue-Qin L Yu-Xiu T Feng W Hong-Liang Z et al QTL mappingand correlations between leaf water potential and drought resistance in rice underupland and lowland environments Acta Agron Sin 200834(2)198ndash206
Ye XDWu XLZhaoH Frehner MNoumlsberger J Potrykus Iet al Altered fructan accumulationin transgenic Lolium multi 1047298orum plants expressing a Bacillus subtilis sacB gene Plant Cell
Rep 200120205ndash
12Zaharieva M Gaulin E Havaux M Acevedo E Monneveux P Drought and heat responses in
the wild wheat relative Aegilops geniculata Roth potential interest for wheatimprovement Crop Sci 2001411321ndash9
Zhang J Zheng HGAartiA Pantuwan G NguyenTT Tripathy JNet alLocatinggenomicregions associated with components of drought resistance in rice comparativemapping within and across species Theor Appl Genet 200110319ndash29
Zhang GH Su Q An LJ Wu S Characterization and expression of a vacuolar Na+H+
antiporter gene from the monocot halophyte Aeluropus littoralis Plant PhysiolBiochem 200846117ndash26
Zhao J QTLs for oil content and their relationships to other agronomic traits in anEuropeantimesChinese oilseed rape population Germany Diss Grorg-Agust Univer-sity of Goettingen 2002
Zhao J Ren W Zhi D Wang L Xia G Arabidopsis DREB1ACBF3 bestowed transgenic tallfescue increased tolerance to drought stress Plant Cell Rep 2007261521ndash8
Zhou W Li Y Zhao BC Ge RC Shen YZ Wang G Huang ZJ Over-expression of TaSTRG gene improves salt and drought tolerance in rice J Plant Physiol 200915(166)1660ndash70
and cost-effective Molecular mapping and analysis of QTL have been
carried out for a number of qualitative and quantitative traits including
stress tolerance which has undoubtedly resulted in a great magnitude
of knowledge and better understanding of the causal genetic phenom-
ena that regulate these traits However limited success has resulted by
using this knowledge to manipulate genes in an effective way for the
improvement of a speci1047297c trait in crops although some crop cultivars
developed through using these tools have performed extremely well
under1047297
eld stressconditions However there are a variety of reasons forthe limited success in terms of achieving enhanced drought tolerance
using the marker-assisted breeding approach For example a major
dif 1047297culty confronting the scientists is the challenge of precise QTL
identi1047297cation In addition a substantial genetictimes environment interac-
tion inconsistent repeatability large number of genes regulating yield
and invalid use of mapping populations have hampered the pursuits
involving mapping of QTL for enhanced drought stress tolerance
Despite these other factors also impede the application of QTL for
genetic improvement of a trait For example due to unfavorable
epistatic interaction it is hard to transfer the effects of a desired allele to
an elite background material (Podlich et al 2004 Collins et al 2008)
Furthermore in some cases QTL from a speci1047297c background do not
show signi1047297cant effects or cease completely in different backgrounds
even under analogous growth conditions (Cho and Hong 2006 Collins
et al 2008) This makes the utilization of QTL more intricate
Transformation of the knowledge acquired from QTL-oriented molec-
ular studies into a well-de1047297ned upshot for the stakeholders is one of the
key challenges confronting the breeders Certainly a multidisciplinary
approach including more speci1047297cally the identi1047297cation of QTL mediated
signal transduction in response to stresses needs to be adopted
Furthermore although QTL cloning procedures are unwieldy they can
lead to a meaningful outcome because a cloned QTL can offer a reliable
markerfor MABas well as furnish an outlinefor thedetection of superior
allelic variants in crop species (Till et al 2007)
Genetic engineering (Transgenic approach) offers a promise whereby
one can expect a substantial improvement in a desired trait within the
shortest stretch of time Engineering genes encoding organic osmolytes
plant growth regulators antioxidants late embryogenesis abundant pro-
teins and transcription factors has resulted into transgenic lines whichhave out-performed under controlled stress conditions In most cases the
claims made by the researchers in terms of performance of transgenic
lines tested under controlled conditions are considerably in1047298ated
However with the exception of a very few notable cases most of the
transgenic lines of differentcrops have been rarely1047297eld-tested Thushow
far these transgenic lines perform under 1047297eld stress conditions is not
known because under natural1047297eld conditions a linecultivar has to face a
multitude of environmental factors other than the drought stress Thus
appraisal of performance of a transgenic line under natural 1047297eld con-
ditionsis mandatory forthe stakeholdersperspective It is also imperative
to note that most of the drought tolerant transgenic lines have been
developed using a single gene transformation which may not be as
productive as if it had been developed using transformation of many
genes Thus transferring a number of prominent genes effectively in-volved in stress tolerance to transgenic plants seems to be a logical
approach Although a large number of genes appear to be involved in
stress tolerance and most of them have been fully characterized the
function of many of them in the mechanism of stress tolerance is yet to be
investigated
References
Abebe T Guenzi AC Martin B Cushman JC Tolerance of mannitol-accumulatingtransgenic wheat to water stress and salinity Plant Physiol 20031311748ndash55
Ashraf M Breeding for salinity tolerance in plants Crit Rev Plant Sci 19941317 ndash42Ashraf M Akram NA Improving salinity tolerance of plants through conventional
breeding and genetic engineering an analytical comparison Biotechnol Adv200927744ndash52 doi101016jbiotechadv200905026
Ashraf M Foolad MR Roles of glycinebetaine and proline in improving plant abioticstress resistance Environ Exp Bot 200759206ndash16
Ashraf M Athar HR Harris PJC Kwon TR Some prospective strategies for improvingcrop salt tolerance Adv Agron 20089745-110
Asins MJ Present and future of quantitative trait locus analysis in plant breeding PlantBreed 2002121281ndash91
Babu RC Nguyen BD Chamarerk V Shanmugasundaram P Chezhian P Jeyaprakash P et alGeneticanalysis of droughtresistance in riceby molecular markers association betweensecondary traits and 1047297eld performance Crop Sci 2003431457ndash69
Babu RC Zhang J Blum A Ho THD Wu R Nguyen HT HVA1 a LEA gene from barleyconfers dehydration tolerance in transgenic rice( Oryzasativa L)via cell membraneprotection Plant Sci 2004166855ndash62
Badawi GH Kawano N Yamauchi Y Over-expression of ascorbate peroxidase in
tobacco chloroplasts enhances the tolerance to salt stress and water de1047297cit PhysiolPlant 2004121231ndash8Badu-Apraku B Yallou CG Registration of striga-resistant and drought tolerant tropical
early maize populations TZE-W Pop DT STR C4 and TZE-Y Pop DT STR C4 J PlantRegistr 20093(1)86ndash90
Badu-Apraku B Menkir A Kling JG Fakorede MAB Registration of 16 striga resistantearly maturing tropical maize inbred lines Crop Sci 2006461410ndash1
Baenziger PS Beecher B Graybosch RA Ibrahim AMH Baltensperger DD Nelson LA et alRegistration of lsquoNEO1643rsquo wheat J Plant Registr 20082(1)36ndash42
Baumlnziger M Setimela PS Hodson D Vivek B Breeding for improved drought tolerance inmaize adapted to southern Africa Proceedings of the 4th International Crop ScienceCongress Brisbane Australia Published on CDROM 2004 26 Sep ndash 1 Oct
Bartels D Sunkar R Drought and salt tolerance in plants Crit Rev Plant Sci 20052423 ndash58Baum M Grandol S Backes G Jahoor A Sabbagh A Ceccarelli S QTLs for agronomic
traits in the Mediterranean environment identi1047297ed in recombinant inbred lines of the cross lsquoArtarsquo H spontaneum 41-1 Theor Appl Genet 20031071215ndash25
Bergman JW Riveland NR Flynn CR Carlson GR Wichman DM Registration of lsquoMorlinrsquo
saf 1047298ower Crop Sci 2001411640Bernier J Kumar A Venuprasad R Spaner D Atlin G A large-effect QTL for
grain yield under reproductive-stage drought stress in upland rice Crop Sci200747507ndash18
BernierJ Kumar A SerrajR SpanerD Atlin G Review breeding uplandrice fordroughtresistance J Sci Food Agric 200888927ndash39
Bernier J Serraj R Kumar A Venuprasad R Impa S Gowdaa RPV et al The large-effectdrought-resistance QTL qtl121 increases water uptake in upland rice Field CropsRes 200911039ndash46
Bhatnagar-Mathur P ReddyDS Lavanya M Yamaguchi-Shinozaki K Sharma KK Stress-inducible expression of Arabidopsis thaliana DREB1A in transgenic peanut ( Arachishypogaea L) increases transpiration ef 1047297ciency under water-limiting conditionsPlant Cell Rep 2007262071ndash82
Bhatnagar-Mathur P Devi MJ Vadez V Sharma KK Differential antioxidative responses intransgenic peanut bear no relationship to their superior transpiration ef 1047297ciency underdrought stress J Plant Physiol 2009166(11)1207ndash17 doi101016jjplph200901001
Bidinger FR Serraj R Rizvi SMH Howarth C Yadav RS Hash CT Field evaluation of droughttolerance QTL effects on phenotype and adaptation in pearl millet [ Pennisetum glaucum(L) R Br] topcross hybrids Field Crops Res 200594(1)14ndash32
Bidinger FR Nepolean T Hash CT Yadav RS Howarth CJ Identi1047297cation of QTLs for grainyield of pearl millet (Pennisetum glaucum (L) R Br) in environments with variablemoisture during grain 1047297lling Crop Sci 200747969ndash80
Bowers JE Abbey C Anderson S Chang C Draye X A high-density geneticrecombination map of sequence-tagged sites for Sorghum as a framework forcomparative structural and evolutionary genomics of tropical grains and grassesGenetics 2003165367ndash86
Brick MA Ogg JB Singh SP Schwartz HF Johnson JJ Pastor-Corrales MA Registration of drought-tolerant rust-resistant high-yielding pinto bean germplasm lineCO46348 J Plant Registr 20082(2)120ndash4
Browne J Tunnacliffe A Burnell A Anhydrobiosis-plant desiccation gene found in anematode Nature 200241638
Busk PK Pages M Regulation of abscisic acid induced transcription Plant Mol Biol199837425ndash35
Carena MJ Wanner DW Development of genetically broad-based inbred lines of maizefor early-maturing (70-80RM) hybrids J Plant Registr 20093107ndash11
Cash SDBrucknerPL Wichman DMKephart KD Berg JEBoynerR et alRegistration of Willow Creek forage wheat J Plant Registr 20093(2)185ndash90
Cattivelli L Rizza F Badeck FW Mazzucotelli E Francia AMEM Mare AT et al Droughttolerance improvement in crop plants an integrated view from breeding to genomics
Field Crops Res 20081051-14Chen M Wang QY Cheng XG Xu ZS Li LC Ye XG et al GmDREB2 a soybean DRE-
binding transcription factor conferred drought and high-salt tolerance intransgenic plants Biochem Biophys Res Commun 2007353299ndash305
Cheng Z Targolli J Huang X Wu R Wheat LEA genes PMA80 and PMA1959 enhancedehydration tolerance of transgenic rice (Oryza sativa L) Mol Breed 20021071ndash82
ChoEK Hong ChB Over-expression of tobacco NtHSP70-1 contributes to drought-stresstolerance in plants Plant Cell Reports 200625349ndash58
Close TJ Dehydrins a commonality in the response of plants to dehydration and lowtemperature Physiol Plant 1997100291ndash6
Collins NC Tardieu F Tuberosa R Quantitative trait loci and crop performance underabiotic stress where do we stand Plant Physiol 2008147469ndash86
Concept Note Combining breeding and biotechnology to develop water ef 1047297cient maizefor Africa (WEMA) Afr Agric Technol Foundation
Courtois BShen L Petalcorin WCarandang S MauleonR LiZ Locating QTLs controllingconstitutive root traits in the rice population IAC 165-Co39 Euphytica 2003134335ndash45
Da-hong L Hui L Yang YL Ping-ping Z Jian-sheng L Down-regulated expression of RACK1geneby RNAinterference enhancesdroughttolerancein rice RiceSci 200916(1)14ndash20
Dalal M Tayal D Chinnusamy V Bansala KC Abiotic stress and ABA-inducible group 4 LEAfrom Brassicanapus plays a key rolein saltand droughttolerance J Biotechnol 2009139137ndash45
Ding Z Li S An X Liu X Qin H Wang D Transgenic expression of MYB15 confers enhancedsensitivity to abscisic acid and improved drought tolerance in Arabidopsis thaliana
J Genet Genomics 20093617ndash29Dubouzet JG Sakuma Y Ito Y Kasuga M Dubouzet EG Miura S et al OsDREB genes in
rice Oryza sativa L encode transcription activators that function in drought high-salt- and cold-responsive gene expression Plant J 200333751ndash63
El-Far IA Allan AY Responses of some wheat cultivars to sowing methods and droughtat different stages of growth Assuit J Agric Sci 199526(1)267ndash77
Eltayeb AE Kawano N Badawi GH Kaminaka H Sanekata T Shibahara T et alOverexpression of monodehydroascorbate reductase in transgenic tobacco confersenhanced tolerance to ozonesalt and polyethyleneglycol stresses Planta 2007225(5)1255ndash64
Falconer DS Introduction to quantitative genetics London New York Longman 1989Feng-ling FU Zhi-Lei F Shi-bing G Shu-feng Z Wan-chen L Evaluation and quantitative
inheritanceof several drought-relativetraits in maize Agric SciChina 20087(3)280ndash90Finkelstein R Gampala S Rock C Abscisic acid signaling in seeds and seedlings Plant
Cell 20021415ndash45Flowers TJ Improving crop salt tolerance J Exp Bot 200455307ndash19Fujita Y Fujita M Satoh R Maruyama K Parvez MM Seki M et al AREB1 is a transcription
Giraudat J Parcy F Bertauche N Gosti F Leung J Morris PC et al Current advances inabscisic acid action and signalling Plant Mol Biol 1994261557ndash77
Gorbalenya AE Koonin EV Helicases amino acid sequence comparisons and structurendashfunction relationships Curr Opin Struct Biol 19933419ndash29
Gosal SS Wani SH Kang MS Biotechnology and drought tolerance J Crop Improvement20092319ndash54
Gubis J Vaňkovaacute R Červenaacute V Draguacuteňovaacute M Hudcovicovaacute M Lichtnerovaacute H et alTransformedtobacco plants with increased toleranceto droughtSouthAfr J Bot200773505ndash11
GuoO Zhang J GaoQ Xing SLi F Wang WDrought tolerancethrough over-expressionof mono ubiquitin in transgenic tobacco J Plant Physiol 20081651745 ndash55
Haley SD Johnson JJ Peairs FB Quick JS Stromberger JA Clayshulte SR et al Registration of lsquoRipperrsquo wheat J Plant Registr 200711ndash6
Han SE Park SR Kwon HB Yi BY Lee GB Byun MO Genetic engineering of drought-resistant tobacco plants by introducingthe trehalose phosphorylase (TP) gene fromPleurotus sajor-caju Plant Cell Tissue Organ Cult 200582151ndash8
Harris K Klein R Mullet J Sorghum stay-green QTL individually reduces post-1047298oweringdrought-induced leaf senescence J Exp Bot 200758327ndash38
Hong BS Zong-Suo L Ming-An S LEA proteins in higher plants structure functiongene expression and regulation Colloids Surf B Biointerf 200545131ndash5
Howarth CJ Yadav RS Successful marker assisted selection for drought tolerance anddisease resistance in pearl milletIGER Innovations 2002
Humphreys MO Humphreys MW Breeding for stress resistance general principles InAshraf M Harris PJC editors Abiotic stresses plant resistance through breedingand molecular approaches 2005 p 19ndash46
Ingram J Bartels D The molecular basis of dehydration tolerance in plants Ann RevPlant Physiol Plant Mol Biol 199647377ndash403
Jaglo-Ottosen KR Gilmour SJ Zarka DG Schabenberger O Thomashow MF ArabidopsisCBF1 overexpression induces COR genes and enhances freezing tolerance Science1998280104ndash6
Jang IC Oh SJ Seo JS Choi WB Song SI Kim CH et al Expression of a bifunctional fusion of the Escherichia coli genes for trehalose-6-phosphate synthase and trehalose-6-phosphate phosphatase in transgenic rice plants increases trehalose accumulationandabiotic stresstolerancewithoutstuntinggrowth Plant Physiol2003131516ndash24
Jeanneau M Gerentes D Foueillassar X Zivy M Vidal J Toppan A et al Improvement of drought tolerance in maize towards the functional validation of the Zm-Asr1 gene andincrease of water use ef 1047297ciency by over-expressing C4-PEPC Biochimie 2002841127ndash35
Jenson KB Palazzo AJ Waldron BL Bushman BS Registration of lsquoFirstStrikersquo Slenderwheatgrass J Plant Registr 2007124ndash5
Joung-youn K Hyung-in C Min-young I Soo-young K Arabidopsis basic leucine zipperproteins that mediate stress-responsive abscisic acid signaling Plant Cell 20024343ndash57
Juskiw PE Helm JH Oro M Nyachiro JM Salmon DF Registration of lsquoBentleyrsquo barley
J Plant Registr 20093(2)119ndash23Kalamaki MS Alexandrou D Lazari D Merkouropoulos G Fotopoulos V Pateraki I et al
Over-expression of a tomato N-acetyl-L -glutamate synthase gene (SlNAGS1) in Arabidopsis thaliana results in high ornithine levels and increased tolerance in saltand drought stresses J Exp Bot 200960(6)1859ndash71
Kamoshita A Babu CR Boopathi NM Fukai S Phenotypic and genotypic analysis of drought-resistance traits for development of rice cultivars adapted to rainfedenvironments Field Crops Res 2008109(103)1-23 doi101016jfcr200806010
KarakasB Ozias-AkinsP Stushnoff C SuefferheldM Rieger M Salinityand drought toleranceof mannitol-accumulating transgenic tobacco Plant Cell Environ 199720609ndash16
Karim S Aronsson H Ericson H Pirhonen M Leyman B Welin B et al Improved droughttolerance without undesired side effects in transgenic plants producing trehalose PlantMol Biol 200764371ndash86
Kasuga M Liu Q Miura S Yamaguchi-Shinozaki K Shinozaki K Improving plant droughtsaltand freezing tolerance by gene transfer of a single stress-inducible transcription factorNat Biotechnol 199917287ndash91
Kindiger M Gaub H Hasegawac M Katsurab Y Ueyamad K Gotob S et al Registrationof lsquoNanryorsquo tall fescue Crop Sci 2006461815ndash6
Kong I Dong J HartGE Characteristics linkage mappositionsand allelic differentiationof Sorghum bicolour (L) Moench by DNA simple-sequence repeats (SSRs) TheorAppl Genet 2000101438ndash48
Kumar R Venuprasad R Atlin GN Genetic analysis of rainfed lowland rice droughttolerance under naturally-occurring stress in eastern India heritability and QTL effects Field Crops Res 200710342ndash52
La1047297tte HR Price AH Courtois B Yield response to water de1047297cit in an upland ricemapping population associations among traits and genetic markers Theor ApplGenet 20041091237ndash46
Lal S Gulyani V Khurana P Overexpressionof HVA1 gene frombarley generates tolerance tosalinity and water stress in transgenic mulberry (Morus indica) Transgenic Res 200817
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63Lanceras J Pantuwan G Jongdee B Toojinda T Quantitative trait loci associated withdrought tolerance at reproductive stage in rice Plant Physiol 2004135384ndash99
Levi A Ovnat L Paterson AH Saranga Y Photosynthesis of cotton near-isogenic linesintrogressed with QTLs for productivity and drought related traits Plant Sci 2009a17788ndash96
Levi A Paterson AH Barak V Yakir D Wang B Chee PW et al Field evaluation of cottonnear-isogenic lines introgressed with QTLs for productivity and drought relatedtraits Mol Breed 2009b23179ndash95
Liu Q Kasuga M Sakuma Y Abe H Miura S Yamaguchi- Shinozaki K et al Twotranscription factors DREB1 and DREB2 with an EREBPAP2 DNA binding domainseparate two cellular signal transduction pathways in drought-and low-temper-ature-responsive gene expression respectively in Arabidopsis Plant Cell 1998101391ndash406
Liu X Hua X Guo J Qi D Wang L Liu Z et al Enhanced tolerance to drought stress intransgenic tobacco plants overexpressing Biotechnol Lett 2008301275ndash80
Liu X Wanga Z Wanga L Wua R Phillips J Deng X LEA 4 group genes from theresurrection plant Boea hygrometrica confer dehydration tolerance in transgenictobacco Plant Sci 200917690ndash8
Luchi S Kobayashi M Taji T Naramoto M Seki M Kato T et al Regulation of droughttolerance by gene manipulation of 9-cis-epoxycarotenoid dioxygenase a keyenzyme in abscisic acid biosynthesis in Arabidopsis Plant J 200127325ndash33
Luo Y Liu YB Dong YX Gao XQ Zhang YS Expression of a putative alfalfa helicase increasestolerance to abioticstress in Arabidopsis by enhancing the capacities for ROS scavengingand osmotic adjustment J Plant Physiol 2009166385ndash94
MacLean JLDaweDC Hardy B Hettel GPRiceAlmanac sourcebook for themostimportanteconomic activity on Earth3rd ed Wallingford England CABI Publishing 2002
Mansour MMF Nitrogen containing compounds and adaptation of plants to salinitystress Biol Plant 200043491ndash500
McKersie BD Bowley SR Harjanto E Leprince O Water-de1047297cit tolerance and 1047297eldperformance of transgenic alfalfa overexpressing superoxide dismutase Plant Physiol19961111177ndash81
McKersie BD Murnaghan J Bowley SR Manipulating freezing tolerance in transgenicplants Acta Physiol Plant 199719485ndash95
Miranda JA Avonce N Suaacuterez R Thevelein JM Dijck PV Iturriaga G A bifunctional TPS-TPP enzyme from yeast confers tolerance to multiple and extreme abiotic-stressconditions in transgenic Arabidopsis Planta 2007226(6)1411ndash21
Mohammadi M Taleei A Zeinali H Naghavi MR Ceccarelli S Grando Baum M QTL analysis for phenologic traits in doubled haploid population of barley Int J AgricBiol 20057(5)820ndash3
Mohanty A Kathuria H Ferjani A Sakamoto A Mohanty P Murata N et al Transgenicsof an elite indica rice variety Pusa Basmati 1 harbouring the codA gene are highlytolerant to salt stress Theor Appl Genet 200210651ndash7
Noaman MM El Sayad AA Asaad FA El Sherbini AM El Bawab AO El Moselhi MA et alRegistration of lsquoGiza 126rsquo barley Crop Sci 199535(6)1710
Noaman MM Ahmed IA El-Sayed AA Abo-El-Enin RA El-Gamal AS El-Sherbiny AM et alRegistration of lsquoGiza 2000rsquo drought-tolerant six-rowed barley for rainfed and newreclaimed areas in Egypt Crop Sci 200747440
Obert DE Evans CP Wesenberg DM Windes JM Erickson CA Jackson EW et alRegistration of lsquoLenetahrsquo spring barley J Plant Registr 20082(2)85ndash7
Oh SJ Jeong JS Kim EH Yi NR Yi SI Jang IC et al Matrix attachment region fromthe chicken lysozyme locus reduces variability in transgene expression andconfers copy number-dependence in transgenic rice plants Plant Cell Rep20054145ndash54
Oh SJ Kim YS Kwon C Park HK Jeong JS Kim JK Overexpression of the transcriptionfactor AP37 in rice improves grain yield under drought conditions Plant Physiol
20092191ndash
200 doi101104pp109137554Owttrim GW RNA helicases and abiotic stress Nucleic AcidsRes 200634(11)3220ndash30Park BJ Liu Z Kanno A Kameya T Increased tolerance to salt and water de1047297cit stress in
transgenic lettuce (Lactuca sativa L) by constitutive expression of LEA PlantGrowth Regul 2005a45165ndash71
Park BJ Liu Z Kanno A Kameya T Genetic improvement of Chinese cabbage for salt anddroughttolerance by constitutiveexpression of a B napus LEAgenePlantSci 2005b169553ndash8
Pastori GM Foyer CH Common componentsnetworks and pathways of crosstoleranceto stress The central role of ldquoredoxrdquo and abscisic acid-mediated controls PlantPhysiol 2002129460ndash8
PellegrineschiA ReynoldsM PachecoM Brito RMAlmeraya R Yamaguchi-ShinozakiK et alStress-induced expression in wheat of the Arabidopsis thaliana DREB1Agenedelayswaterstress symptoms under greenhouse conditions Genome 200447493ndash500
Perl A Perl-Treves R Galili S Aviv D Shalgi E Malkin S et al Enhanced oxidative-stressdefense in transgenic potato overexpressing tomato Cu Zn superoxide dismutaseTheor Appl Genet 199385568ndash76
Podlich DW Winkler CR Cooper M Mapping as you go an effective approach formarker-assisted selection of complex traits Crop Sci 2004441560ndash71
182 M Ashraf Biotechnology Advances 28 (2010) 169ndash183
Pratt RC Casey MA Registration of maize germplasm line Oh605 Crop Sci 2006461004ndash5Qin F Kakimoto M Sakuma Y Maruyama K Osakabe Y Tran LSP et al Regulation and
functional analysis of ZmDREB2A in response to drought and heat stress in Zea mays LPlant J 20075054ndash69
Quan R Shang M Zhang H Zhao Y Zhang J Engineering of enhanced glycinebetainesynthesis improves drought tolerance in maize Plant Biotechnol J 20042477ndash86
Quarrie SA Gulli M Calestani C Steed A Marmiroli N Location of a gene regulatingdrought-induced abscisic acid production on the long arm of chromosome 5A of wheat Theor Appl Genet 199489794ndash800
QuickJS Stromberger JA Clayshulte S Clifford B Johnson JJ Peairs FB et al Registrationof lsquoPrairie Redrsquo wheat Crop Sci 2001411362ndash3
RajaramS Prospects andpromise ofwheatbreedingin the21stcentury Euphytica 20011193-15Rajaram S Role of conventional plant breeding and biotechnology in future wheat
production Turk J Agric Forest 200529105ndash11Rami JF Dufour P Trouche G Fliedel G Mestres C Davrieux F et al Quantitative trait
loci for grain quality productivity morphological and agronomical traits insorghum (Sorghum bicolor L Moench) Theor Appl Genet 199897605ndash16
Rassaa N Salahb HBH Latiri K Thermal responses of durum wheat Triticum durum toearly water stress consequence on leaf and 1047298ower development Plant Biol Pathol2008331(5)363ndash71
Reddy LJ Nigam SN Rao RCN Reddy NS Registration of ICGV 87354 peanut germplasmwith drought tolerance and rust resistance Crop Sci 200141274ndash5
Ribaut JM Ragot M Marker-assisted selection to improve drought adaptation in maize thebackcross approach perspectives limitations and alternatives J Exp Bot 200658351ndash60
Robin S Pathan MS Courtois B La1047297tte R Carandang S Lanceras S et al Mappingosmotic adjustment in an advanced back-cross inbred population of rice Theor
Appl Genet 2003107(7)1288ndash96Romero C Belles JM Vaya JL Serrano R Culianez-Macia FA Expression of the yeast
trehalose-6-phosphate synthase gene in transgenic tobacco plants pleiotropicphenotypes include drought tolerance Planta 1997201293ndash7
Ronde JAD Cress WA Krugerd GHJ Strasserd RJ Van Staden J Photosynthetic responseof transgenic soybean plants containing an Arabidopsis P5CR gene during heat anddrought stress J Plant Physiol 20041611211ndash24
Salvi S Tuberosa R To clone or not to clone plant QTLs present and future challengesTrends Plant Sci 200510297ndash304
Sanchez AC Subudhi PK Rosenow DT Nguyen HT Mapping QTLs associated with droughtresistance in sorghum (Sorghum bicolor L Moench) Plant Mol Biol 200248713ndash26
Saranga Y Menz M Jiang CX Wright RJ Yakir D Paterson AH Genomic dissection of genotypetimesenvironment interactions conferring adaptation of cotton to aridconditions Genome Res 2001111988ndash95
Sari-Gorla M Krajewski P Di Fonzo N Villa M Frova C Genetic analysis of droughttolerance in maize by molecular markers II Plant height and 1047298owering Theor ApplGenet 199999289ndash95
Seki M Narusaka M Ishida J Nanjo T Fujita M Oono Y et al Monitoring the expressionpro1047297les of 7000 Arabidopsis genes under drought cold and high-salinity stressesusing a full-length cDNA microarray Plant J 200231279ndash92
Serraj R Sinclair TR Osmolyte accumulation can it really increase crop yield underdrought conditions Plant Cell Environ 200225333ndash41
Serraj R Krishnamurthy L Kashiwagi J Kumar J Chandra S Crouch JH Variation in roottraits of chickpea (Cicer arietinum L) grown under terminal drought Field CropsRes 200488115ndash27
Serraj R Hash CT Rizvi MHS Sharma A Yadav RS Bidinger FR Recent advances in marker-assisted selection for drought tolerance in pearl millet Plant Prod Sci 20058(3)334ndash7
Shen YG Du BX Zhang WK Zhang JS Chen SY AhCMO regulated by stresses in Atriplexhortensis can improve drought tolerance in transgenic tobacco Theor Appl Genet2002105815ndash21
Shinozaki K Yamaguchi-Shinozaki K Molecular responses to dehydration and lowtemperature differences and cross-talk between two stress signaling pathwaysCurr Opin Plant Biol 20003217ndash23
Singh KB Omar M Saxena MC Johansen C Registration of FLIP 87-59C a drought-tolerant chickpea germplasm line Crop Sci 199636(2)1ndash2
Singh SP Teran H Gutierrez JA Registration of SEA 5 and SEA 13 drought tolerant drybean germplasm Crop Sci 200141276ndash7
Singh SP Teran H Lema M Schwartz HF Miklas PN Registration of white moldresistant dry bean germplasm line A 195 J Plant Registr 2007162ndash3
Sivamani E Bahieldin A Wraith JM Improved biomass productivity and water useef 1047297ciency under water de1047297cit conditions in transgenic wheat constitutivelyexpressing the barley HVA1 gene Plant Sci 20001551ndash9
Skovmand B Reynolds MP DeLacy IH Searching genetic resources for physiologicaltraits with potential for increasing yield In Reynolds MP Ortiz-Monasterio IMcNab A editors Application of Physiology in Wheat Breeding 2001 p 17ndash28
Steele K Novel upland rice variety bred using marker-assisted selection and client-oriented breeding released in Jharkhand India Bangor University 2009
Steele KA Price AH Shashidar HE Witcombe JR Marker-assistedselection to introgressrice QTLs controlling root traitsinto an Indianupland rice variety Theor Appl Genet2006112208ndash21
Steele KA Virk DS Kumar R Prasad SC Witcombe JR Field evaluation of upland ricelines selected for QTLs controlling root traits Field Crops Res 2007101180 ndash6
Sunkar R Kapoor A Zhu JK Post transcriptional induction of two CuZn superoxidedismutase genes in Arabidopsis is mediated by down regulation of miR398 andimportant for oxidative stress tolerance Plant Cell 2006182051ndash65
Talame V Sanguineti MC Chiapparino E Bahri H Ben Salem M Forster BP et alIdenti1047297cation of Hordeum spontaneum QTL alleles improving 1047297eld performance of barley grown under rainfed conditions Ann Appl Bot 2004144309ndash20
Tanner NKCordinO BanroquesJ DoereM Linder PThe Q Motif a newlyidenti1047297ed motif in DEAD box helicases may regulate ATP binding and hydrolysis Mol Cell 200311127ndash38
Taramino G Tarchini R Ferrario S Lee M Pe ME Characterization and mapping of simplesequence repeats (SSRs) in Sorghum bicolor Theor Appl Genet 19979566ndash72
Teulat B Monneveux P Wery J Borriegraves C Souyris I Charrier A et al Relationshipsbetween relative water content andgrowth parameters in barleya QTLstudy NewPhytol 199713799-107
Thi Lang N Chi Buu B Fine mapping for drought tolerance in rice ( Oryza sativa L)Omonrice 2008169-15Till BJ Comai L Henikoff S Tillering and ecotillering for crop improvement In
Varshney RK Tuberosa R editors Genomics-assisted crop improvement genomicsapproaches and platforms vol 1 2007 p 333ndash50
Tondelli A Francia E Barabaschi D Aprile A Skinner JS Stockinger EJ et al Mappingregulatory genes as candidates for cold and drought stress tolerance in barleyTheor Appl Genet 2006112445ndash54
Tuberosa R Salvi S Genomics approaches to improve drought tolerance in cropsTrends Plant Sci 200611405ndash12
Valkoun JJ Wheat pre-breeding using wild progenitors Euphytica 200111917ndash23Vashisht AA Tuteja N Stress responsive DEAD-box helicases a new pathway to
engineer plant stress tolerance J Photochem Photobiol B Biol 200684150 ndash60Vendruscolo ECG Schuster I Pileggi M Scapim CA Molinari HBC Marur CJ et al Stress-
induced synthesis of proline confers tolerance to water de1047297cit in transgenic wheat J Plant Physiol 20071641367ndash76
Vienne D Leonardi A Damerval C Zivy M Genetics of proteome variation for QTL characterization application to drought stress responses in maize J Exp Bot 199950303ndash9
Villareal RL Mujeeb-Kazi A Rajaram S Toro ED Morphological variability in somesynthetic hexaploid wheats derived from Triticum turgidum times T tauschii J GenetBreed 1994487-16
Vinh NT Paterson AH Genome mapping and its implication for stress resistance inplants In Ashraf M Harris PJC editors Abiotic stresses plant resistance throughbreeding and molecular approaches 2005
Vinocur B Altman A Recent advances in engineering plant tolerance to abiotic stressachievements and limitations Curr Opin Biotechnol 200516123ndash32
Wang YJ Hao YJ Zhang ZG Chen T Zhang JS Chen SY Isolation of trehalose-6-phosphate phosphatase gene from tobacco and its functional analysis in yeast cells
J Plant Physiol 2005162215ndash23Wu R Garg A Engineering rice plants with trehalose-producing genes improves
tolerance to drought salt and low temperature ISB News Report 2003Xiao B Huang Y Tang N Xiong L Overexpression of LEA gene in rice improves drought
resistance under 1047297eld conditions Theor Appl Genet 200711535ndash46Xinglai P Sangang X Qiannying P Yinhong S Registration of lsquo Jinmai 50rsquo wheat Crop Sci
200646983ndash5Xiong L Lee H Ishitani M Zhu JK Regulation of osmotic stress-responsive gene
expression by the LOS6ABA1 locus in Arabidopsis J Biol Chem 20022778588ndash96Xu GW Magill CW Shertz KF Hart GE A RFLP linkage map of Sorghum bicolor (L)
Moench Theor Appl Genet 199489139ndash45Xu D Duan X Wang B Hong BHo THD Wu RExpression of a late embryogenesis abundant
protein gene HVA1 from barley confers tolerance to water de1047297cit and salt stress intransgenic rice Plant Physiol 1996110249ndash57
Yadav RS Hash CT Bidinger FR Devos KM Howarth CJ Genomic regions associatedwith grain yield and aspects of post-1047298owering drought tolerance in pearl milletacross stress environments and testers background Euphytica 2004136265ndash77
YamadaM MorishitaH Urano K Shiozaki N Yamaguchi-Shinozaki K ShinozakiK et alEffects of free proline accumulation in petunias under drought stress J Exp Bot2005561975ndash81
Yang WJ Rich PJ Axtell JD Wood KV Bonham CC Ejeta G et al Genotypic variation forglycinebetaine in sorghum Crop Sci 200343162ndash9
Yan-Ying QU Ping MU Xue-Qin L Yu-Xiu T Feng W Hong-Liang Z et al QTL mappingand correlations between leaf water potential and drought resistance in rice underupland and lowland environments Acta Agron Sin 200834(2)198ndash206
Ye XDWu XLZhaoH Frehner MNoumlsberger J Potrykus Iet al Altered fructan accumulationin transgenic Lolium multi 1047298orum plants expressing a Bacillus subtilis sacB gene Plant Cell
Rep 200120205ndash
12Zaharieva M Gaulin E Havaux M Acevedo E Monneveux P Drought and heat responses in
the wild wheat relative Aegilops geniculata Roth potential interest for wheatimprovement Crop Sci 2001411321ndash9
Zhang J Zheng HGAartiA Pantuwan G NguyenTT Tripathy JNet alLocatinggenomicregions associated with components of drought resistance in rice comparativemapping within and across species Theor Appl Genet 200110319ndash29
Zhang GH Su Q An LJ Wu S Characterization and expression of a vacuolar Na+H+
antiporter gene from the monocot halophyte Aeluropus littoralis Plant PhysiolBiochem 200846117ndash26
Zhao J QTLs for oil content and their relationships to other agronomic traits in anEuropeantimesChinese oilseed rape population Germany Diss Grorg-Agust Univer-sity of Goettingen 2002
Zhao J Ren W Zhi D Wang L Xia G Arabidopsis DREB1ACBF3 bestowed transgenic tallfescue increased tolerance to drought stress Plant Cell Rep 2007261521ndash8
Zhou W Li Y Zhao BC Ge RC Shen YZ Wang G Huang ZJ Over-expression of TaSTRG gene improves salt and drought tolerance in rice J Plant Physiol 200915(166)1660ndash70
and cost-effective Molecular mapping and analysis of QTL have been
carried out for a number of qualitative and quantitative traits including
stress tolerance which has undoubtedly resulted in a great magnitude
of knowledge and better understanding of the causal genetic phenom-
ena that regulate these traits However limited success has resulted by
using this knowledge to manipulate genes in an effective way for the
improvement of a speci1047297c trait in crops although some crop cultivars
developed through using these tools have performed extremely well
under1047297
eld stressconditions However there are a variety of reasons forthe limited success in terms of achieving enhanced drought tolerance
using the marker-assisted breeding approach For example a major
dif 1047297culty confronting the scientists is the challenge of precise QTL
identi1047297cation In addition a substantial genetictimes environment interac-
tion inconsistent repeatability large number of genes regulating yield
and invalid use of mapping populations have hampered the pursuits
involving mapping of QTL for enhanced drought stress tolerance
Despite these other factors also impede the application of QTL for
genetic improvement of a trait For example due to unfavorable
epistatic interaction it is hard to transfer the effects of a desired allele to
an elite background material (Podlich et al 2004 Collins et al 2008)
Furthermore in some cases QTL from a speci1047297c background do not
show signi1047297cant effects or cease completely in different backgrounds
even under analogous growth conditions (Cho and Hong 2006 Collins
et al 2008) This makes the utilization of QTL more intricate
Transformation of the knowledge acquired from QTL-oriented molec-
ular studies into a well-de1047297ned upshot for the stakeholders is one of the
key challenges confronting the breeders Certainly a multidisciplinary
approach including more speci1047297cally the identi1047297cation of QTL mediated
signal transduction in response to stresses needs to be adopted
Furthermore although QTL cloning procedures are unwieldy they can
lead to a meaningful outcome because a cloned QTL can offer a reliable
markerfor MABas well as furnish an outlinefor thedetection of superior
allelic variants in crop species (Till et al 2007)
Genetic engineering (Transgenic approach) offers a promise whereby
one can expect a substantial improvement in a desired trait within the
shortest stretch of time Engineering genes encoding organic osmolytes
plant growth regulators antioxidants late embryogenesis abundant pro-
teins and transcription factors has resulted into transgenic lines whichhave out-performed under controlled stress conditions In most cases the
claims made by the researchers in terms of performance of transgenic
lines tested under controlled conditions are considerably in1047298ated
However with the exception of a very few notable cases most of the
transgenic lines of differentcrops have been rarely1047297eld-tested Thushow
far these transgenic lines perform under 1047297eld stress conditions is not
known because under natural1047297eld conditions a linecultivar has to face a
multitude of environmental factors other than the drought stress Thus
appraisal of performance of a transgenic line under natural 1047297eld con-
ditionsis mandatory forthe stakeholdersperspective It is also imperative
to note that most of the drought tolerant transgenic lines have been
developed using a single gene transformation which may not be as
productive as if it had been developed using transformation of many
genes Thus transferring a number of prominent genes effectively in-volved in stress tolerance to transgenic plants seems to be a logical
approach Although a large number of genes appear to be involved in
stress tolerance and most of them have been fully characterized the
function of many of them in the mechanism of stress tolerance is yet to be
investigated
References
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Ashraf M Breeding for salinity tolerance in plants Crit Rev Plant Sci 19941317 ndash42Ashraf M Akram NA Improving salinity tolerance of plants through conventional
breeding and genetic engineering an analytical comparison Biotechnol Adv200927744ndash52 doi101016jbiotechadv200905026
Ashraf M Foolad MR Roles of glycinebetaine and proline in improving plant abioticstress resistance Environ Exp Bot 200759206ndash16
Ashraf M Athar HR Harris PJC Kwon TR Some prospective strategies for improvingcrop salt tolerance Adv Agron 20089745-110
Asins MJ Present and future of quantitative trait locus analysis in plant breeding PlantBreed 2002121281ndash91
Babu RC Nguyen BD Chamarerk V Shanmugasundaram P Chezhian P Jeyaprakash P et alGeneticanalysis of droughtresistance in riceby molecular markers association betweensecondary traits and 1047297eld performance Crop Sci 2003431457ndash69
Babu RC Zhang J Blum A Ho THD Wu R Nguyen HT HVA1 a LEA gene from barleyconfers dehydration tolerance in transgenic rice( Oryzasativa L)via cell membraneprotection Plant Sci 2004166855ndash62
Badawi GH Kawano N Yamauchi Y Over-expression of ascorbate peroxidase in
tobacco chloroplasts enhances the tolerance to salt stress and water de1047297cit PhysiolPlant 2004121231ndash8Badu-Apraku B Yallou CG Registration of striga-resistant and drought tolerant tropical
early maize populations TZE-W Pop DT STR C4 and TZE-Y Pop DT STR C4 J PlantRegistr 20093(1)86ndash90
Badu-Apraku B Menkir A Kling JG Fakorede MAB Registration of 16 striga resistantearly maturing tropical maize inbred lines Crop Sci 2006461410ndash1
Baenziger PS Beecher B Graybosch RA Ibrahim AMH Baltensperger DD Nelson LA et alRegistration of lsquoNEO1643rsquo wheat J Plant Registr 20082(1)36ndash42
Baumlnziger M Setimela PS Hodson D Vivek B Breeding for improved drought tolerance inmaize adapted to southern Africa Proceedings of the 4th International Crop ScienceCongress Brisbane Australia Published on CDROM 2004 26 Sep ndash 1 Oct
Bartels D Sunkar R Drought and salt tolerance in plants Crit Rev Plant Sci 20052423 ndash58Baum M Grandol S Backes G Jahoor A Sabbagh A Ceccarelli S QTLs for agronomic
traits in the Mediterranean environment identi1047297ed in recombinant inbred lines of the cross lsquoArtarsquo H spontaneum 41-1 Theor Appl Genet 20031071215ndash25
Bergman JW Riveland NR Flynn CR Carlson GR Wichman DM Registration of lsquoMorlinrsquo
saf 1047298ower Crop Sci 2001411640Bernier J Kumar A Venuprasad R Spaner D Atlin G A large-effect QTL for
grain yield under reproductive-stage drought stress in upland rice Crop Sci200747507ndash18
BernierJ Kumar A SerrajR SpanerD Atlin G Review breeding uplandrice fordroughtresistance J Sci Food Agric 200888927ndash39
Bernier J Serraj R Kumar A Venuprasad R Impa S Gowdaa RPV et al The large-effectdrought-resistance QTL qtl121 increases water uptake in upland rice Field CropsRes 200911039ndash46
Bhatnagar-Mathur P ReddyDS Lavanya M Yamaguchi-Shinozaki K Sharma KK Stress-inducible expression of Arabidopsis thaliana DREB1A in transgenic peanut ( Arachishypogaea L) increases transpiration ef 1047297ciency under water-limiting conditionsPlant Cell Rep 2007262071ndash82
Bhatnagar-Mathur P Devi MJ Vadez V Sharma KK Differential antioxidative responses intransgenic peanut bear no relationship to their superior transpiration ef 1047297ciency underdrought stress J Plant Physiol 2009166(11)1207ndash17 doi101016jjplph200901001
Bidinger FR Serraj R Rizvi SMH Howarth C Yadav RS Hash CT Field evaluation of droughttolerance QTL effects on phenotype and adaptation in pearl millet [ Pennisetum glaucum(L) R Br] topcross hybrids Field Crops Res 200594(1)14ndash32
Bidinger FR Nepolean T Hash CT Yadav RS Howarth CJ Identi1047297cation of QTLs for grainyield of pearl millet (Pennisetum glaucum (L) R Br) in environments with variablemoisture during grain 1047297lling Crop Sci 200747969ndash80
Bowers JE Abbey C Anderson S Chang C Draye X A high-density geneticrecombination map of sequence-tagged sites for Sorghum as a framework forcomparative structural and evolutionary genomics of tropical grains and grassesGenetics 2003165367ndash86
Brick MA Ogg JB Singh SP Schwartz HF Johnson JJ Pastor-Corrales MA Registration of drought-tolerant rust-resistant high-yielding pinto bean germplasm lineCO46348 J Plant Registr 20082(2)120ndash4
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Busk PK Pages M Regulation of abscisic acid induced transcription Plant Mol Biol199837425ndash35
Carena MJ Wanner DW Development of genetically broad-based inbred lines of maizefor early-maturing (70-80RM) hybrids J Plant Registr 20093107ndash11
Cash SDBrucknerPL Wichman DMKephart KD Berg JEBoynerR et alRegistration of Willow Creek forage wheat J Plant Registr 20093(2)185ndash90
Cattivelli L Rizza F Badeck FW Mazzucotelli E Francia AMEM Mare AT et al Droughttolerance improvement in crop plants an integrated view from breeding to genomics
Field Crops Res 20081051-14Chen M Wang QY Cheng XG Xu ZS Li LC Ye XG et al GmDREB2 a soybean DRE-
binding transcription factor conferred drought and high-salt tolerance intransgenic plants Biochem Biophys Res Commun 2007353299ndash305
Cheng Z Targolli J Huang X Wu R Wheat LEA genes PMA80 and PMA1959 enhancedehydration tolerance of transgenic rice (Oryza sativa L) Mol Breed 20021071ndash82
ChoEK Hong ChB Over-expression of tobacco NtHSP70-1 contributes to drought-stresstolerance in plants Plant Cell Reports 200625349ndash58
Close TJ Dehydrins a commonality in the response of plants to dehydration and lowtemperature Physiol Plant 1997100291ndash6
Collins NC Tardieu F Tuberosa R Quantitative trait loci and crop performance underabiotic stress where do we stand Plant Physiol 2008147469ndash86
Concept Note Combining breeding and biotechnology to develop water ef 1047297cient maizefor Africa (WEMA) Afr Agric Technol Foundation
Courtois BShen L Petalcorin WCarandang S MauleonR LiZ Locating QTLs controllingconstitutive root traits in the rice population IAC 165-Co39 Euphytica 2003134335ndash45
Da-hong L Hui L Yang YL Ping-ping Z Jian-sheng L Down-regulated expression of RACK1geneby RNAinterference enhancesdroughttolerancein rice RiceSci 200916(1)14ndash20
Dalal M Tayal D Chinnusamy V Bansala KC Abiotic stress and ABA-inducible group 4 LEAfrom Brassicanapus plays a key rolein saltand droughttolerance J Biotechnol 2009139137ndash45
Ding Z Li S An X Liu X Qin H Wang D Transgenic expression of MYB15 confers enhancedsensitivity to abscisic acid and improved drought tolerance in Arabidopsis thaliana
J Genet Genomics 20093617ndash29Dubouzet JG Sakuma Y Ito Y Kasuga M Dubouzet EG Miura S et al OsDREB genes in
rice Oryza sativa L encode transcription activators that function in drought high-salt- and cold-responsive gene expression Plant J 200333751ndash63
El-Far IA Allan AY Responses of some wheat cultivars to sowing methods and droughtat different stages of growth Assuit J Agric Sci 199526(1)267ndash77
Eltayeb AE Kawano N Badawi GH Kaminaka H Sanekata T Shibahara T et alOverexpression of monodehydroascorbate reductase in transgenic tobacco confersenhanced tolerance to ozonesalt and polyethyleneglycol stresses Planta 2007225(5)1255ndash64
Falconer DS Introduction to quantitative genetics London New York Longman 1989Feng-ling FU Zhi-Lei F Shi-bing G Shu-feng Z Wan-chen L Evaluation and quantitative
inheritanceof several drought-relativetraits in maize Agric SciChina 20087(3)280ndash90Finkelstein R Gampala S Rock C Abscisic acid signaling in seeds and seedlings Plant
Cell 20021415ndash45Flowers TJ Improving crop salt tolerance J Exp Bot 200455307ndash19Fujita Y Fujita M Satoh R Maruyama K Parvez MM Seki M et al AREB1 is a transcription
Giraudat J Parcy F Bertauche N Gosti F Leung J Morris PC et al Current advances inabscisic acid action and signalling Plant Mol Biol 1994261557ndash77
Gorbalenya AE Koonin EV Helicases amino acid sequence comparisons and structurendashfunction relationships Curr Opin Struct Biol 19933419ndash29
Gosal SS Wani SH Kang MS Biotechnology and drought tolerance J Crop Improvement20092319ndash54
Gubis J Vaňkovaacute R Červenaacute V Draguacuteňovaacute M Hudcovicovaacute M Lichtnerovaacute H et alTransformedtobacco plants with increased toleranceto droughtSouthAfr J Bot200773505ndash11
GuoO Zhang J GaoQ Xing SLi F Wang WDrought tolerancethrough over-expressionof mono ubiquitin in transgenic tobacco J Plant Physiol 20081651745 ndash55
Haley SD Johnson JJ Peairs FB Quick JS Stromberger JA Clayshulte SR et al Registration of lsquoRipperrsquo wheat J Plant Registr 200711ndash6
Han SE Park SR Kwon HB Yi BY Lee GB Byun MO Genetic engineering of drought-resistant tobacco plants by introducingthe trehalose phosphorylase (TP) gene fromPleurotus sajor-caju Plant Cell Tissue Organ Cult 200582151ndash8
Harris K Klein R Mullet J Sorghum stay-green QTL individually reduces post-1047298oweringdrought-induced leaf senescence J Exp Bot 200758327ndash38
Hong BS Zong-Suo L Ming-An S LEA proteins in higher plants structure functiongene expression and regulation Colloids Surf B Biointerf 200545131ndash5
Howarth CJ Yadav RS Successful marker assisted selection for drought tolerance anddisease resistance in pearl milletIGER Innovations 2002
Humphreys MO Humphreys MW Breeding for stress resistance general principles InAshraf M Harris PJC editors Abiotic stresses plant resistance through breedingand molecular approaches 2005 p 19ndash46
Ingram J Bartels D The molecular basis of dehydration tolerance in plants Ann RevPlant Physiol Plant Mol Biol 199647377ndash403
Jaglo-Ottosen KR Gilmour SJ Zarka DG Schabenberger O Thomashow MF ArabidopsisCBF1 overexpression induces COR genes and enhances freezing tolerance Science1998280104ndash6
Jang IC Oh SJ Seo JS Choi WB Song SI Kim CH et al Expression of a bifunctional fusion of the Escherichia coli genes for trehalose-6-phosphate synthase and trehalose-6-phosphate phosphatase in transgenic rice plants increases trehalose accumulationandabiotic stresstolerancewithoutstuntinggrowth Plant Physiol2003131516ndash24
Jeanneau M Gerentes D Foueillassar X Zivy M Vidal J Toppan A et al Improvement of drought tolerance in maize towards the functional validation of the Zm-Asr1 gene andincrease of water use ef 1047297ciency by over-expressing C4-PEPC Biochimie 2002841127ndash35
Jenson KB Palazzo AJ Waldron BL Bushman BS Registration of lsquoFirstStrikersquo Slenderwheatgrass J Plant Registr 2007124ndash5
Joung-youn K Hyung-in C Min-young I Soo-young K Arabidopsis basic leucine zipperproteins that mediate stress-responsive abscisic acid signaling Plant Cell 20024343ndash57
Juskiw PE Helm JH Oro M Nyachiro JM Salmon DF Registration of lsquoBentleyrsquo barley
J Plant Registr 20093(2)119ndash23Kalamaki MS Alexandrou D Lazari D Merkouropoulos G Fotopoulos V Pateraki I et al
Over-expression of a tomato N-acetyl-L -glutamate synthase gene (SlNAGS1) in Arabidopsis thaliana results in high ornithine levels and increased tolerance in saltand drought stresses J Exp Bot 200960(6)1859ndash71
Kamoshita A Babu CR Boopathi NM Fukai S Phenotypic and genotypic analysis of drought-resistance traits for development of rice cultivars adapted to rainfedenvironments Field Crops Res 2008109(103)1-23 doi101016jfcr200806010
KarakasB Ozias-AkinsP Stushnoff C SuefferheldM Rieger M Salinityand drought toleranceof mannitol-accumulating transgenic tobacco Plant Cell Environ 199720609ndash16
Karim S Aronsson H Ericson H Pirhonen M Leyman B Welin B et al Improved droughttolerance without undesired side effects in transgenic plants producing trehalose PlantMol Biol 200764371ndash86
Kasuga M Liu Q Miura S Yamaguchi-Shinozaki K Shinozaki K Improving plant droughtsaltand freezing tolerance by gene transfer of a single stress-inducible transcription factorNat Biotechnol 199917287ndash91
Kindiger M Gaub H Hasegawac M Katsurab Y Ueyamad K Gotob S et al Registrationof lsquoNanryorsquo tall fescue Crop Sci 2006461815ndash6
Kong I Dong J HartGE Characteristics linkage mappositionsand allelic differentiationof Sorghum bicolour (L) Moench by DNA simple-sequence repeats (SSRs) TheorAppl Genet 2000101438ndash48
Kumar R Venuprasad R Atlin GN Genetic analysis of rainfed lowland rice droughttolerance under naturally-occurring stress in eastern India heritability and QTL effects Field Crops Res 200710342ndash52
La1047297tte HR Price AH Courtois B Yield response to water de1047297cit in an upland ricemapping population associations among traits and genetic markers Theor ApplGenet 20041091237ndash46
Lal S Gulyani V Khurana P Overexpressionof HVA1 gene frombarley generates tolerance tosalinity and water stress in transgenic mulberry (Morus indica) Transgenic Res 200817
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63Lanceras J Pantuwan G Jongdee B Toojinda T Quantitative trait loci associated withdrought tolerance at reproductive stage in rice Plant Physiol 2004135384ndash99
Levi A Ovnat L Paterson AH Saranga Y Photosynthesis of cotton near-isogenic linesintrogressed with QTLs for productivity and drought related traits Plant Sci 2009a17788ndash96
Levi A Paterson AH Barak V Yakir D Wang B Chee PW et al Field evaluation of cottonnear-isogenic lines introgressed with QTLs for productivity and drought relatedtraits Mol Breed 2009b23179ndash95
Liu Q Kasuga M Sakuma Y Abe H Miura S Yamaguchi- Shinozaki K et al Twotranscription factors DREB1 and DREB2 with an EREBPAP2 DNA binding domainseparate two cellular signal transduction pathways in drought-and low-temper-ature-responsive gene expression respectively in Arabidopsis Plant Cell 1998101391ndash406
Liu X Hua X Guo J Qi D Wang L Liu Z et al Enhanced tolerance to drought stress intransgenic tobacco plants overexpressing Biotechnol Lett 2008301275ndash80
Liu X Wanga Z Wanga L Wua R Phillips J Deng X LEA 4 group genes from theresurrection plant Boea hygrometrica confer dehydration tolerance in transgenictobacco Plant Sci 200917690ndash8
Luchi S Kobayashi M Taji T Naramoto M Seki M Kato T et al Regulation of droughttolerance by gene manipulation of 9-cis-epoxycarotenoid dioxygenase a keyenzyme in abscisic acid biosynthesis in Arabidopsis Plant J 200127325ndash33
Luo Y Liu YB Dong YX Gao XQ Zhang YS Expression of a putative alfalfa helicase increasestolerance to abioticstress in Arabidopsis by enhancing the capacities for ROS scavengingand osmotic adjustment J Plant Physiol 2009166385ndash94
MacLean JLDaweDC Hardy B Hettel GPRiceAlmanac sourcebook for themostimportanteconomic activity on Earth3rd ed Wallingford England CABI Publishing 2002
Mansour MMF Nitrogen containing compounds and adaptation of plants to salinitystress Biol Plant 200043491ndash500
McKersie BD Bowley SR Harjanto E Leprince O Water-de1047297cit tolerance and 1047297eldperformance of transgenic alfalfa overexpressing superoxide dismutase Plant Physiol19961111177ndash81
McKersie BD Murnaghan J Bowley SR Manipulating freezing tolerance in transgenicplants Acta Physiol Plant 199719485ndash95
Miranda JA Avonce N Suaacuterez R Thevelein JM Dijck PV Iturriaga G A bifunctional TPS-TPP enzyme from yeast confers tolerance to multiple and extreme abiotic-stressconditions in transgenic Arabidopsis Planta 2007226(6)1411ndash21
Mohammadi M Taleei A Zeinali H Naghavi MR Ceccarelli S Grando Baum M QTL analysis for phenologic traits in doubled haploid population of barley Int J AgricBiol 20057(5)820ndash3
Mohanty A Kathuria H Ferjani A Sakamoto A Mohanty P Murata N et al Transgenicsof an elite indica rice variety Pusa Basmati 1 harbouring the codA gene are highlytolerant to salt stress Theor Appl Genet 200210651ndash7
Noaman MM El Sayad AA Asaad FA El Sherbini AM El Bawab AO El Moselhi MA et alRegistration of lsquoGiza 126rsquo barley Crop Sci 199535(6)1710
Noaman MM Ahmed IA El-Sayed AA Abo-El-Enin RA El-Gamal AS El-Sherbiny AM et alRegistration of lsquoGiza 2000rsquo drought-tolerant six-rowed barley for rainfed and newreclaimed areas in Egypt Crop Sci 200747440
Obert DE Evans CP Wesenberg DM Windes JM Erickson CA Jackson EW et alRegistration of lsquoLenetahrsquo spring barley J Plant Registr 20082(2)85ndash7
Oh SJ Jeong JS Kim EH Yi NR Yi SI Jang IC et al Matrix attachment region fromthe chicken lysozyme locus reduces variability in transgene expression andconfers copy number-dependence in transgenic rice plants Plant Cell Rep20054145ndash54
Oh SJ Kim YS Kwon C Park HK Jeong JS Kim JK Overexpression of the transcriptionfactor AP37 in rice improves grain yield under drought conditions Plant Physiol
20092191ndash
200 doi101104pp109137554Owttrim GW RNA helicases and abiotic stress Nucleic AcidsRes 200634(11)3220ndash30Park BJ Liu Z Kanno A Kameya T Increased tolerance to salt and water de1047297cit stress in
transgenic lettuce (Lactuca sativa L) by constitutive expression of LEA PlantGrowth Regul 2005a45165ndash71
Park BJ Liu Z Kanno A Kameya T Genetic improvement of Chinese cabbage for salt anddroughttolerance by constitutiveexpression of a B napus LEAgenePlantSci 2005b169553ndash8
Pastori GM Foyer CH Common componentsnetworks and pathways of crosstoleranceto stress The central role of ldquoredoxrdquo and abscisic acid-mediated controls PlantPhysiol 2002129460ndash8
PellegrineschiA ReynoldsM PachecoM Brito RMAlmeraya R Yamaguchi-ShinozakiK et alStress-induced expression in wheat of the Arabidopsis thaliana DREB1Agenedelayswaterstress symptoms under greenhouse conditions Genome 200447493ndash500
Perl A Perl-Treves R Galili S Aviv D Shalgi E Malkin S et al Enhanced oxidative-stressdefense in transgenic potato overexpressing tomato Cu Zn superoxide dismutaseTheor Appl Genet 199385568ndash76
Podlich DW Winkler CR Cooper M Mapping as you go an effective approach formarker-assisted selection of complex traits Crop Sci 2004441560ndash71
182 M Ashraf Biotechnology Advances 28 (2010) 169ndash183
Pratt RC Casey MA Registration of maize germplasm line Oh605 Crop Sci 2006461004ndash5Qin F Kakimoto M Sakuma Y Maruyama K Osakabe Y Tran LSP et al Regulation and
functional analysis of ZmDREB2A in response to drought and heat stress in Zea mays LPlant J 20075054ndash69
Quan R Shang M Zhang H Zhao Y Zhang J Engineering of enhanced glycinebetainesynthesis improves drought tolerance in maize Plant Biotechnol J 20042477ndash86
Quarrie SA Gulli M Calestani C Steed A Marmiroli N Location of a gene regulatingdrought-induced abscisic acid production on the long arm of chromosome 5A of wheat Theor Appl Genet 199489794ndash800
QuickJS Stromberger JA Clayshulte S Clifford B Johnson JJ Peairs FB et al Registrationof lsquoPrairie Redrsquo wheat Crop Sci 2001411362ndash3
RajaramS Prospects andpromise ofwheatbreedingin the21stcentury Euphytica 20011193-15Rajaram S Role of conventional plant breeding and biotechnology in future wheat
production Turk J Agric Forest 200529105ndash11Rami JF Dufour P Trouche G Fliedel G Mestres C Davrieux F et al Quantitative trait
loci for grain quality productivity morphological and agronomical traits insorghum (Sorghum bicolor L Moench) Theor Appl Genet 199897605ndash16
Rassaa N Salahb HBH Latiri K Thermal responses of durum wheat Triticum durum toearly water stress consequence on leaf and 1047298ower development Plant Biol Pathol2008331(5)363ndash71
Reddy LJ Nigam SN Rao RCN Reddy NS Registration of ICGV 87354 peanut germplasmwith drought tolerance and rust resistance Crop Sci 200141274ndash5
Ribaut JM Ragot M Marker-assisted selection to improve drought adaptation in maize thebackcross approach perspectives limitations and alternatives J Exp Bot 200658351ndash60
Robin S Pathan MS Courtois B La1047297tte R Carandang S Lanceras S et al Mappingosmotic adjustment in an advanced back-cross inbred population of rice Theor
Appl Genet 2003107(7)1288ndash96Romero C Belles JM Vaya JL Serrano R Culianez-Macia FA Expression of the yeast
trehalose-6-phosphate synthase gene in transgenic tobacco plants pleiotropicphenotypes include drought tolerance Planta 1997201293ndash7
Ronde JAD Cress WA Krugerd GHJ Strasserd RJ Van Staden J Photosynthetic responseof transgenic soybean plants containing an Arabidopsis P5CR gene during heat anddrought stress J Plant Physiol 20041611211ndash24
Salvi S Tuberosa R To clone or not to clone plant QTLs present and future challengesTrends Plant Sci 200510297ndash304
Sanchez AC Subudhi PK Rosenow DT Nguyen HT Mapping QTLs associated with droughtresistance in sorghum (Sorghum bicolor L Moench) Plant Mol Biol 200248713ndash26
Saranga Y Menz M Jiang CX Wright RJ Yakir D Paterson AH Genomic dissection of genotypetimesenvironment interactions conferring adaptation of cotton to aridconditions Genome Res 2001111988ndash95
Sari-Gorla M Krajewski P Di Fonzo N Villa M Frova C Genetic analysis of droughttolerance in maize by molecular markers II Plant height and 1047298owering Theor ApplGenet 199999289ndash95
Seki M Narusaka M Ishida J Nanjo T Fujita M Oono Y et al Monitoring the expressionpro1047297les of 7000 Arabidopsis genes under drought cold and high-salinity stressesusing a full-length cDNA microarray Plant J 200231279ndash92
Serraj R Sinclair TR Osmolyte accumulation can it really increase crop yield underdrought conditions Plant Cell Environ 200225333ndash41
Serraj R Krishnamurthy L Kashiwagi J Kumar J Chandra S Crouch JH Variation in roottraits of chickpea (Cicer arietinum L) grown under terminal drought Field CropsRes 200488115ndash27
Serraj R Hash CT Rizvi MHS Sharma A Yadav RS Bidinger FR Recent advances in marker-assisted selection for drought tolerance in pearl millet Plant Prod Sci 20058(3)334ndash7
Shen YG Du BX Zhang WK Zhang JS Chen SY AhCMO regulated by stresses in Atriplexhortensis can improve drought tolerance in transgenic tobacco Theor Appl Genet2002105815ndash21
Shinozaki K Yamaguchi-Shinozaki K Molecular responses to dehydration and lowtemperature differences and cross-talk between two stress signaling pathwaysCurr Opin Plant Biol 20003217ndash23
Singh KB Omar M Saxena MC Johansen C Registration of FLIP 87-59C a drought-tolerant chickpea germplasm line Crop Sci 199636(2)1ndash2
Singh SP Teran H Gutierrez JA Registration of SEA 5 and SEA 13 drought tolerant drybean germplasm Crop Sci 200141276ndash7
Singh SP Teran H Lema M Schwartz HF Miklas PN Registration of white moldresistant dry bean germplasm line A 195 J Plant Registr 2007162ndash3
Sivamani E Bahieldin A Wraith JM Improved biomass productivity and water useef 1047297ciency under water de1047297cit conditions in transgenic wheat constitutivelyexpressing the barley HVA1 gene Plant Sci 20001551ndash9
Skovmand B Reynolds MP DeLacy IH Searching genetic resources for physiologicaltraits with potential for increasing yield In Reynolds MP Ortiz-Monasterio IMcNab A editors Application of Physiology in Wheat Breeding 2001 p 17ndash28
Steele K Novel upland rice variety bred using marker-assisted selection and client-oriented breeding released in Jharkhand India Bangor University 2009
Steele KA Price AH Shashidar HE Witcombe JR Marker-assistedselection to introgressrice QTLs controlling root traitsinto an Indianupland rice variety Theor Appl Genet2006112208ndash21
Steele KA Virk DS Kumar R Prasad SC Witcombe JR Field evaluation of upland ricelines selected for QTLs controlling root traits Field Crops Res 2007101180 ndash6
Sunkar R Kapoor A Zhu JK Post transcriptional induction of two CuZn superoxidedismutase genes in Arabidopsis is mediated by down regulation of miR398 andimportant for oxidative stress tolerance Plant Cell 2006182051ndash65
Talame V Sanguineti MC Chiapparino E Bahri H Ben Salem M Forster BP et alIdenti1047297cation of Hordeum spontaneum QTL alleles improving 1047297eld performance of barley grown under rainfed conditions Ann Appl Bot 2004144309ndash20
Tanner NKCordinO BanroquesJ DoereM Linder PThe Q Motif a newlyidenti1047297ed motif in DEAD box helicases may regulate ATP binding and hydrolysis Mol Cell 200311127ndash38
Taramino G Tarchini R Ferrario S Lee M Pe ME Characterization and mapping of simplesequence repeats (SSRs) in Sorghum bicolor Theor Appl Genet 19979566ndash72
Teulat B Monneveux P Wery J Borriegraves C Souyris I Charrier A et al Relationshipsbetween relative water content andgrowth parameters in barleya QTLstudy NewPhytol 199713799-107
Thi Lang N Chi Buu B Fine mapping for drought tolerance in rice ( Oryza sativa L)Omonrice 2008169-15Till BJ Comai L Henikoff S Tillering and ecotillering for crop improvement In
Varshney RK Tuberosa R editors Genomics-assisted crop improvement genomicsapproaches and platforms vol 1 2007 p 333ndash50
Tondelli A Francia E Barabaschi D Aprile A Skinner JS Stockinger EJ et al Mappingregulatory genes as candidates for cold and drought stress tolerance in barleyTheor Appl Genet 2006112445ndash54
Tuberosa R Salvi S Genomics approaches to improve drought tolerance in cropsTrends Plant Sci 200611405ndash12
Valkoun JJ Wheat pre-breeding using wild progenitors Euphytica 200111917ndash23Vashisht AA Tuteja N Stress responsive DEAD-box helicases a new pathway to
engineer plant stress tolerance J Photochem Photobiol B Biol 200684150 ndash60Vendruscolo ECG Schuster I Pileggi M Scapim CA Molinari HBC Marur CJ et al Stress-
induced synthesis of proline confers tolerance to water de1047297cit in transgenic wheat J Plant Physiol 20071641367ndash76
Vienne D Leonardi A Damerval C Zivy M Genetics of proteome variation for QTL characterization application to drought stress responses in maize J Exp Bot 199950303ndash9
Villareal RL Mujeeb-Kazi A Rajaram S Toro ED Morphological variability in somesynthetic hexaploid wheats derived from Triticum turgidum times T tauschii J GenetBreed 1994487-16
Vinh NT Paterson AH Genome mapping and its implication for stress resistance inplants In Ashraf M Harris PJC editors Abiotic stresses plant resistance throughbreeding and molecular approaches 2005
Vinocur B Altman A Recent advances in engineering plant tolerance to abiotic stressachievements and limitations Curr Opin Biotechnol 200516123ndash32
Wang YJ Hao YJ Zhang ZG Chen T Zhang JS Chen SY Isolation of trehalose-6-phosphate phosphatase gene from tobacco and its functional analysis in yeast cells
J Plant Physiol 2005162215ndash23Wu R Garg A Engineering rice plants with trehalose-producing genes improves
tolerance to drought salt and low temperature ISB News Report 2003Xiao B Huang Y Tang N Xiong L Overexpression of LEA gene in rice improves drought
resistance under 1047297eld conditions Theor Appl Genet 200711535ndash46Xinglai P Sangang X Qiannying P Yinhong S Registration of lsquo Jinmai 50rsquo wheat Crop Sci
200646983ndash5Xiong L Lee H Ishitani M Zhu JK Regulation of osmotic stress-responsive gene
expression by the LOS6ABA1 locus in Arabidopsis J Biol Chem 20022778588ndash96Xu GW Magill CW Shertz KF Hart GE A RFLP linkage map of Sorghum bicolor (L)
Moench Theor Appl Genet 199489139ndash45Xu D Duan X Wang B Hong BHo THD Wu RExpression of a late embryogenesis abundant
protein gene HVA1 from barley confers tolerance to water de1047297cit and salt stress intransgenic rice Plant Physiol 1996110249ndash57
Yadav RS Hash CT Bidinger FR Devos KM Howarth CJ Genomic regions associatedwith grain yield and aspects of post-1047298owering drought tolerance in pearl milletacross stress environments and testers background Euphytica 2004136265ndash77
YamadaM MorishitaH Urano K Shiozaki N Yamaguchi-Shinozaki K ShinozakiK et alEffects of free proline accumulation in petunias under drought stress J Exp Bot2005561975ndash81
Yang WJ Rich PJ Axtell JD Wood KV Bonham CC Ejeta G et al Genotypic variation forglycinebetaine in sorghum Crop Sci 200343162ndash9
Yan-Ying QU Ping MU Xue-Qin L Yu-Xiu T Feng W Hong-Liang Z et al QTL mappingand correlations between leaf water potential and drought resistance in rice underupland and lowland environments Acta Agron Sin 200834(2)198ndash206
Ye XDWu XLZhaoH Frehner MNoumlsberger J Potrykus Iet al Altered fructan accumulationin transgenic Lolium multi 1047298orum plants expressing a Bacillus subtilis sacB gene Plant Cell
Rep 200120205ndash
12Zaharieva M Gaulin E Havaux M Acevedo E Monneveux P Drought and heat responses in
the wild wheat relative Aegilops geniculata Roth potential interest for wheatimprovement Crop Sci 2001411321ndash9
Zhang J Zheng HGAartiA Pantuwan G NguyenTT Tripathy JNet alLocatinggenomicregions associated with components of drought resistance in rice comparativemapping within and across species Theor Appl Genet 200110319ndash29
Zhang GH Su Q An LJ Wu S Characterization and expression of a vacuolar Na+H+
antiporter gene from the monocot halophyte Aeluropus littoralis Plant PhysiolBiochem 200846117ndash26
Zhao J QTLs for oil content and their relationships to other agronomic traits in anEuropeantimesChinese oilseed rape population Germany Diss Grorg-Agust Univer-sity of Goettingen 2002
Zhao J Ren W Zhi D Wang L Xia G Arabidopsis DREB1ACBF3 bestowed transgenic tallfescue increased tolerance to drought stress Plant Cell Rep 2007261521ndash8
Zhou W Li Y Zhao BC Ge RC Shen YZ Wang G Huang ZJ Over-expression of TaSTRG gene improves salt and drought tolerance in rice J Plant Physiol 200915(166)1660ndash70
and cost-effective Molecular mapping and analysis of QTL have been
carried out for a number of qualitative and quantitative traits including
stress tolerance which has undoubtedly resulted in a great magnitude
of knowledge and better understanding of the causal genetic phenom-
ena that regulate these traits However limited success has resulted by
using this knowledge to manipulate genes in an effective way for the
improvement of a speci1047297c trait in crops although some crop cultivars
developed through using these tools have performed extremely well
under1047297
eld stressconditions However there are a variety of reasons forthe limited success in terms of achieving enhanced drought tolerance
using the marker-assisted breeding approach For example a major
dif 1047297culty confronting the scientists is the challenge of precise QTL
identi1047297cation In addition a substantial genetictimes environment interac-
tion inconsistent repeatability large number of genes regulating yield
and invalid use of mapping populations have hampered the pursuits
involving mapping of QTL for enhanced drought stress tolerance
Despite these other factors also impede the application of QTL for
genetic improvement of a trait For example due to unfavorable
epistatic interaction it is hard to transfer the effects of a desired allele to
an elite background material (Podlich et al 2004 Collins et al 2008)
Furthermore in some cases QTL from a speci1047297c background do not
show signi1047297cant effects or cease completely in different backgrounds
even under analogous growth conditions (Cho and Hong 2006 Collins
et al 2008) This makes the utilization of QTL more intricate
Transformation of the knowledge acquired from QTL-oriented molec-
ular studies into a well-de1047297ned upshot for the stakeholders is one of the
key challenges confronting the breeders Certainly a multidisciplinary
approach including more speci1047297cally the identi1047297cation of QTL mediated
signal transduction in response to stresses needs to be adopted
Furthermore although QTL cloning procedures are unwieldy they can
lead to a meaningful outcome because a cloned QTL can offer a reliable
markerfor MABas well as furnish an outlinefor thedetection of superior
allelic variants in crop species (Till et al 2007)
Genetic engineering (Transgenic approach) offers a promise whereby
one can expect a substantial improvement in a desired trait within the
shortest stretch of time Engineering genes encoding organic osmolytes
plant growth regulators antioxidants late embryogenesis abundant pro-
teins and transcription factors has resulted into transgenic lines whichhave out-performed under controlled stress conditions In most cases the
claims made by the researchers in terms of performance of transgenic
lines tested under controlled conditions are considerably in1047298ated
However with the exception of a very few notable cases most of the
transgenic lines of differentcrops have been rarely1047297eld-tested Thushow
far these transgenic lines perform under 1047297eld stress conditions is not
known because under natural1047297eld conditions a linecultivar has to face a
multitude of environmental factors other than the drought stress Thus
appraisal of performance of a transgenic line under natural 1047297eld con-
ditionsis mandatory forthe stakeholdersperspective It is also imperative
to note that most of the drought tolerant transgenic lines have been
developed using a single gene transformation which may not be as
productive as if it had been developed using transformation of many
genes Thus transferring a number of prominent genes effectively in-volved in stress tolerance to transgenic plants seems to be a logical
approach Although a large number of genes appear to be involved in
stress tolerance and most of them have been fully characterized the
function of many of them in the mechanism of stress tolerance is yet to be
investigated
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Ashraf M Breeding for salinity tolerance in plants Crit Rev Plant Sci 19941317 ndash42Ashraf M Akram NA Improving salinity tolerance of plants through conventional
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Ashraf M Foolad MR Roles of glycinebetaine and proline in improving plant abioticstress resistance Environ Exp Bot 200759206ndash16
Ashraf M Athar HR Harris PJC Kwon TR Some prospective strategies for improvingcrop salt tolerance Adv Agron 20089745-110
Asins MJ Present and future of quantitative trait locus analysis in plant breeding PlantBreed 2002121281ndash91
Babu RC Nguyen BD Chamarerk V Shanmugasundaram P Chezhian P Jeyaprakash P et alGeneticanalysis of droughtresistance in riceby molecular markers association betweensecondary traits and 1047297eld performance Crop Sci 2003431457ndash69
Babu RC Zhang J Blum A Ho THD Wu R Nguyen HT HVA1 a LEA gene from barleyconfers dehydration tolerance in transgenic rice( Oryzasativa L)via cell membraneprotection Plant Sci 2004166855ndash62
Badawi GH Kawano N Yamauchi Y Over-expression of ascorbate peroxidase in
tobacco chloroplasts enhances the tolerance to salt stress and water de1047297cit PhysiolPlant 2004121231ndash8Badu-Apraku B Yallou CG Registration of striga-resistant and drought tolerant tropical
early maize populations TZE-W Pop DT STR C4 and TZE-Y Pop DT STR C4 J PlantRegistr 20093(1)86ndash90
Badu-Apraku B Menkir A Kling JG Fakorede MAB Registration of 16 striga resistantearly maturing tropical maize inbred lines Crop Sci 2006461410ndash1
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Baumlnziger M Setimela PS Hodson D Vivek B Breeding for improved drought tolerance inmaize adapted to southern Africa Proceedings of the 4th International Crop ScienceCongress Brisbane Australia Published on CDROM 2004 26 Sep ndash 1 Oct
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traits in the Mediterranean environment identi1047297ed in recombinant inbred lines of the cross lsquoArtarsquo H spontaneum 41-1 Theor Appl Genet 20031071215ndash25
Bergman JW Riveland NR Flynn CR Carlson GR Wichman DM Registration of lsquoMorlinrsquo
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grain yield under reproductive-stage drought stress in upland rice Crop Sci200747507ndash18
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Bhatnagar-Mathur P ReddyDS Lavanya M Yamaguchi-Shinozaki K Sharma KK Stress-inducible expression of Arabidopsis thaliana DREB1A in transgenic peanut ( Arachishypogaea L) increases transpiration ef 1047297ciency under water-limiting conditionsPlant Cell Rep 2007262071ndash82
Bhatnagar-Mathur P Devi MJ Vadez V Sharma KK Differential antioxidative responses intransgenic peanut bear no relationship to their superior transpiration ef 1047297ciency underdrought stress J Plant Physiol 2009166(11)1207ndash17 doi101016jjplph200901001
Bidinger FR Serraj R Rizvi SMH Howarth C Yadav RS Hash CT Field evaluation of droughttolerance QTL effects on phenotype and adaptation in pearl millet [ Pennisetum glaucum(L) R Br] topcross hybrids Field Crops Res 200594(1)14ndash32
Bidinger FR Nepolean T Hash CT Yadav RS Howarth CJ Identi1047297cation of QTLs for grainyield of pearl millet (Pennisetum glaucum (L) R Br) in environments with variablemoisture during grain 1047297lling Crop Sci 200747969ndash80
Bowers JE Abbey C Anderson S Chang C Draye X A high-density geneticrecombination map of sequence-tagged sites for Sorghum as a framework forcomparative structural and evolutionary genomics of tropical grains and grassesGenetics 2003165367ndash86
Brick MA Ogg JB Singh SP Schwartz HF Johnson JJ Pastor-Corrales MA Registration of drought-tolerant rust-resistant high-yielding pinto bean germplasm lineCO46348 J Plant Registr 20082(2)120ndash4
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Busk PK Pages M Regulation of abscisic acid induced transcription Plant Mol Biol199837425ndash35
Carena MJ Wanner DW Development of genetically broad-based inbred lines of maizefor early-maturing (70-80RM) hybrids J Plant Registr 20093107ndash11
Cash SDBrucknerPL Wichman DMKephart KD Berg JEBoynerR et alRegistration of Willow Creek forage wheat J Plant Registr 20093(2)185ndash90
Cattivelli L Rizza F Badeck FW Mazzucotelli E Francia AMEM Mare AT et al Droughttolerance improvement in crop plants an integrated view from breeding to genomics
Field Crops Res 20081051-14Chen M Wang QY Cheng XG Xu ZS Li LC Ye XG et al GmDREB2 a soybean DRE-
binding transcription factor conferred drought and high-salt tolerance intransgenic plants Biochem Biophys Res Commun 2007353299ndash305
Cheng Z Targolli J Huang X Wu R Wheat LEA genes PMA80 and PMA1959 enhancedehydration tolerance of transgenic rice (Oryza sativa L) Mol Breed 20021071ndash82
ChoEK Hong ChB Over-expression of tobacco NtHSP70-1 contributes to drought-stresstolerance in plants Plant Cell Reports 200625349ndash58
Close TJ Dehydrins a commonality in the response of plants to dehydration and lowtemperature Physiol Plant 1997100291ndash6
Collins NC Tardieu F Tuberosa R Quantitative trait loci and crop performance underabiotic stress where do we stand Plant Physiol 2008147469ndash86
Concept Note Combining breeding and biotechnology to develop water ef 1047297cient maizefor Africa (WEMA) Afr Agric Technol Foundation
Courtois BShen L Petalcorin WCarandang S MauleonR LiZ Locating QTLs controllingconstitutive root traits in the rice population IAC 165-Co39 Euphytica 2003134335ndash45
Da-hong L Hui L Yang YL Ping-ping Z Jian-sheng L Down-regulated expression of RACK1geneby RNAinterference enhancesdroughttolerancein rice RiceSci 200916(1)14ndash20
Dalal M Tayal D Chinnusamy V Bansala KC Abiotic stress and ABA-inducible group 4 LEAfrom Brassicanapus plays a key rolein saltand droughttolerance J Biotechnol 2009139137ndash45
Ding Z Li S An X Liu X Qin H Wang D Transgenic expression of MYB15 confers enhancedsensitivity to abscisic acid and improved drought tolerance in Arabidopsis thaliana
J Genet Genomics 20093617ndash29Dubouzet JG Sakuma Y Ito Y Kasuga M Dubouzet EG Miura S et al OsDREB genes in
rice Oryza sativa L encode transcription activators that function in drought high-salt- and cold-responsive gene expression Plant J 200333751ndash63
El-Far IA Allan AY Responses of some wheat cultivars to sowing methods and droughtat different stages of growth Assuit J Agric Sci 199526(1)267ndash77
Eltayeb AE Kawano N Badawi GH Kaminaka H Sanekata T Shibahara T et alOverexpression of monodehydroascorbate reductase in transgenic tobacco confersenhanced tolerance to ozonesalt and polyethyleneglycol stresses Planta 2007225(5)1255ndash64
Falconer DS Introduction to quantitative genetics London New York Longman 1989Feng-ling FU Zhi-Lei F Shi-bing G Shu-feng Z Wan-chen L Evaluation and quantitative
inheritanceof several drought-relativetraits in maize Agric SciChina 20087(3)280ndash90Finkelstein R Gampala S Rock C Abscisic acid signaling in seeds and seedlings Plant
Cell 20021415ndash45Flowers TJ Improving crop salt tolerance J Exp Bot 200455307ndash19Fujita Y Fujita M Satoh R Maruyama K Parvez MM Seki M et al AREB1 is a transcription
Giraudat J Parcy F Bertauche N Gosti F Leung J Morris PC et al Current advances inabscisic acid action and signalling Plant Mol Biol 1994261557ndash77
Gorbalenya AE Koonin EV Helicases amino acid sequence comparisons and structurendashfunction relationships Curr Opin Struct Biol 19933419ndash29
Gosal SS Wani SH Kang MS Biotechnology and drought tolerance J Crop Improvement20092319ndash54
Gubis J Vaňkovaacute R Červenaacute V Draguacuteňovaacute M Hudcovicovaacute M Lichtnerovaacute H et alTransformedtobacco plants with increased toleranceto droughtSouthAfr J Bot200773505ndash11
GuoO Zhang J GaoQ Xing SLi F Wang WDrought tolerancethrough over-expressionof mono ubiquitin in transgenic tobacco J Plant Physiol 20081651745 ndash55
Haley SD Johnson JJ Peairs FB Quick JS Stromberger JA Clayshulte SR et al Registration of lsquoRipperrsquo wheat J Plant Registr 200711ndash6
Han SE Park SR Kwon HB Yi BY Lee GB Byun MO Genetic engineering of drought-resistant tobacco plants by introducingthe trehalose phosphorylase (TP) gene fromPleurotus sajor-caju Plant Cell Tissue Organ Cult 200582151ndash8
Harris K Klein R Mullet J Sorghum stay-green QTL individually reduces post-1047298oweringdrought-induced leaf senescence J Exp Bot 200758327ndash38
Hong BS Zong-Suo L Ming-An S LEA proteins in higher plants structure functiongene expression and regulation Colloids Surf B Biointerf 200545131ndash5
Howarth CJ Yadav RS Successful marker assisted selection for drought tolerance anddisease resistance in pearl milletIGER Innovations 2002
Humphreys MO Humphreys MW Breeding for stress resistance general principles InAshraf M Harris PJC editors Abiotic stresses plant resistance through breedingand molecular approaches 2005 p 19ndash46
Ingram J Bartels D The molecular basis of dehydration tolerance in plants Ann RevPlant Physiol Plant Mol Biol 199647377ndash403
Jaglo-Ottosen KR Gilmour SJ Zarka DG Schabenberger O Thomashow MF ArabidopsisCBF1 overexpression induces COR genes and enhances freezing tolerance Science1998280104ndash6
Jang IC Oh SJ Seo JS Choi WB Song SI Kim CH et al Expression of a bifunctional fusion of the Escherichia coli genes for trehalose-6-phosphate synthase and trehalose-6-phosphate phosphatase in transgenic rice plants increases trehalose accumulationandabiotic stresstolerancewithoutstuntinggrowth Plant Physiol2003131516ndash24
Jeanneau M Gerentes D Foueillassar X Zivy M Vidal J Toppan A et al Improvement of drought tolerance in maize towards the functional validation of the Zm-Asr1 gene andincrease of water use ef 1047297ciency by over-expressing C4-PEPC Biochimie 2002841127ndash35
Jenson KB Palazzo AJ Waldron BL Bushman BS Registration of lsquoFirstStrikersquo Slenderwheatgrass J Plant Registr 2007124ndash5
Joung-youn K Hyung-in C Min-young I Soo-young K Arabidopsis basic leucine zipperproteins that mediate stress-responsive abscisic acid signaling Plant Cell 20024343ndash57
Juskiw PE Helm JH Oro M Nyachiro JM Salmon DF Registration of lsquoBentleyrsquo barley
J Plant Registr 20093(2)119ndash23Kalamaki MS Alexandrou D Lazari D Merkouropoulos G Fotopoulos V Pateraki I et al
Over-expression of a tomato N-acetyl-L -glutamate synthase gene (SlNAGS1) in Arabidopsis thaliana results in high ornithine levels and increased tolerance in saltand drought stresses J Exp Bot 200960(6)1859ndash71
Kamoshita A Babu CR Boopathi NM Fukai S Phenotypic and genotypic analysis of drought-resistance traits for development of rice cultivars adapted to rainfedenvironments Field Crops Res 2008109(103)1-23 doi101016jfcr200806010
KarakasB Ozias-AkinsP Stushnoff C SuefferheldM Rieger M Salinityand drought toleranceof mannitol-accumulating transgenic tobacco Plant Cell Environ 199720609ndash16
Karim S Aronsson H Ericson H Pirhonen M Leyman B Welin B et al Improved droughttolerance without undesired side effects in transgenic plants producing trehalose PlantMol Biol 200764371ndash86
Kasuga M Liu Q Miura S Yamaguchi-Shinozaki K Shinozaki K Improving plant droughtsaltand freezing tolerance by gene transfer of a single stress-inducible transcription factorNat Biotechnol 199917287ndash91
Kindiger M Gaub H Hasegawac M Katsurab Y Ueyamad K Gotob S et al Registrationof lsquoNanryorsquo tall fescue Crop Sci 2006461815ndash6
Kong I Dong J HartGE Characteristics linkage mappositionsand allelic differentiationof Sorghum bicolour (L) Moench by DNA simple-sequence repeats (SSRs) TheorAppl Genet 2000101438ndash48
Kumar R Venuprasad R Atlin GN Genetic analysis of rainfed lowland rice droughttolerance under naturally-occurring stress in eastern India heritability and QTL effects Field Crops Res 200710342ndash52
La1047297tte HR Price AH Courtois B Yield response to water de1047297cit in an upland ricemapping population associations among traits and genetic markers Theor ApplGenet 20041091237ndash46
Lal S Gulyani V Khurana P Overexpressionof HVA1 gene frombarley generates tolerance tosalinity and water stress in transgenic mulberry (Morus indica) Transgenic Res 200817
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63Lanceras J Pantuwan G Jongdee B Toojinda T Quantitative trait loci associated withdrought tolerance at reproductive stage in rice Plant Physiol 2004135384ndash99
Levi A Ovnat L Paterson AH Saranga Y Photosynthesis of cotton near-isogenic linesintrogressed with QTLs for productivity and drought related traits Plant Sci 2009a17788ndash96
Levi A Paterson AH Barak V Yakir D Wang B Chee PW et al Field evaluation of cottonnear-isogenic lines introgressed with QTLs for productivity and drought relatedtraits Mol Breed 2009b23179ndash95
Liu Q Kasuga M Sakuma Y Abe H Miura S Yamaguchi- Shinozaki K et al Twotranscription factors DREB1 and DREB2 with an EREBPAP2 DNA binding domainseparate two cellular signal transduction pathways in drought-and low-temper-ature-responsive gene expression respectively in Arabidopsis Plant Cell 1998101391ndash406
Liu X Hua X Guo J Qi D Wang L Liu Z et al Enhanced tolerance to drought stress intransgenic tobacco plants overexpressing Biotechnol Lett 2008301275ndash80
Liu X Wanga Z Wanga L Wua R Phillips J Deng X LEA 4 group genes from theresurrection plant Boea hygrometrica confer dehydration tolerance in transgenictobacco Plant Sci 200917690ndash8
Luchi S Kobayashi M Taji T Naramoto M Seki M Kato T et al Regulation of droughttolerance by gene manipulation of 9-cis-epoxycarotenoid dioxygenase a keyenzyme in abscisic acid biosynthesis in Arabidopsis Plant J 200127325ndash33
Luo Y Liu YB Dong YX Gao XQ Zhang YS Expression of a putative alfalfa helicase increasestolerance to abioticstress in Arabidopsis by enhancing the capacities for ROS scavengingand osmotic adjustment J Plant Physiol 2009166385ndash94
MacLean JLDaweDC Hardy B Hettel GPRiceAlmanac sourcebook for themostimportanteconomic activity on Earth3rd ed Wallingford England CABI Publishing 2002
Mansour MMF Nitrogen containing compounds and adaptation of plants to salinitystress Biol Plant 200043491ndash500
McKersie BD Bowley SR Harjanto E Leprince O Water-de1047297cit tolerance and 1047297eldperformance of transgenic alfalfa overexpressing superoxide dismutase Plant Physiol19961111177ndash81
McKersie BD Murnaghan J Bowley SR Manipulating freezing tolerance in transgenicplants Acta Physiol Plant 199719485ndash95
Miranda JA Avonce N Suaacuterez R Thevelein JM Dijck PV Iturriaga G A bifunctional TPS-TPP enzyme from yeast confers tolerance to multiple and extreme abiotic-stressconditions in transgenic Arabidopsis Planta 2007226(6)1411ndash21
Mohammadi M Taleei A Zeinali H Naghavi MR Ceccarelli S Grando Baum M QTL analysis for phenologic traits in doubled haploid population of barley Int J AgricBiol 20057(5)820ndash3
Mohanty A Kathuria H Ferjani A Sakamoto A Mohanty P Murata N et al Transgenicsof an elite indica rice variety Pusa Basmati 1 harbouring the codA gene are highlytolerant to salt stress Theor Appl Genet 200210651ndash7
Noaman MM El Sayad AA Asaad FA El Sherbini AM El Bawab AO El Moselhi MA et alRegistration of lsquoGiza 126rsquo barley Crop Sci 199535(6)1710
Noaman MM Ahmed IA El-Sayed AA Abo-El-Enin RA El-Gamal AS El-Sherbiny AM et alRegistration of lsquoGiza 2000rsquo drought-tolerant six-rowed barley for rainfed and newreclaimed areas in Egypt Crop Sci 200747440
Obert DE Evans CP Wesenberg DM Windes JM Erickson CA Jackson EW et alRegistration of lsquoLenetahrsquo spring barley J Plant Registr 20082(2)85ndash7
Oh SJ Jeong JS Kim EH Yi NR Yi SI Jang IC et al Matrix attachment region fromthe chicken lysozyme locus reduces variability in transgene expression andconfers copy number-dependence in transgenic rice plants Plant Cell Rep20054145ndash54
Oh SJ Kim YS Kwon C Park HK Jeong JS Kim JK Overexpression of the transcriptionfactor AP37 in rice improves grain yield under drought conditions Plant Physiol
20092191ndash
200 doi101104pp109137554Owttrim GW RNA helicases and abiotic stress Nucleic AcidsRes 200634(11)3220ndash30Park BJ Liu Z Kanno A Kameya T Increased tolerance to salt and water de1047297cit stress in
transgenic lettuce (Lactuca sativa L) by constitutive expression of LEA PlantGrowth Regul 2005a45165ndash71
Park BJ Liu Z Kanno A Kameya T Genetic improvement of Chinese cabbage for salt anddroughttolerance by constitutiveexpression of a B napus LEAgenePlantSci 2005b169553ndash8
Pastori GM Foyer CH Common componentsnetworks and pathways of crosstoleranceto stress The central role of ldquoredoxrdquo and abscisic acid-mediated controls PlantPhysiol 2002129460ndash8
PellegrineschiA ReynoldsM PachecoM Brito RMAlmeraya R Yamaguchi-ShinozakiK et alStress-induced expression in wheat of the Arabidopsis thaliana DREB1Agenedelayswaterstress symptoms under greenhouse conditions Genome 200447493ndash500
Perl A Perl-Treves R Galili S Aviv D Shalgi E Malkin S et al Enhanced oxidative-stressdefense in transgenic potato overexpressing tomato Cu Zn superoxide dismutaseTheor Appl Genet 199385568ndash76
Podlich DW Winkler CR Cooper M Mapping as you go an effective approach formarker-assisted selection of complex traits Crop Sci 2004441560ndash71
182 M Ashraf Biotechnology Advances 28 (2010) 169ndash183
Pratt RC Casey MA Registration of maize germplasm line Oh605 Crop Sci 2006461004ndash5Qin F Kakimoto M Sakuma Y Maruyama K Osakabe Y Tran LSP et al Regulation and
functional analysis of ZmDREB2A in response to drought and heat stress in Zea mays LPlant J 20075054ndash69
Quan R Shang M Zhang H Zhao Y Zhang J Engineering of enhanced glycinebetainesynthesis improves drought tolerance in maize Plant Biotechnol J 20042477ndash86
Quarrie SA Gulli M Calestani C Steed A Marmiroli N Location of a gene regulatingdrought-induced abscisic acid production on the long arm of chromosome 5A of wheat Theor Appl Genet 199489794ndash800
QuickJS Stromberger JA Clayshulte S Clifford B Johnson JJ Peairs FB et al Registrationof lsquoPrairie Redrsquo wheat Crop Sci 2001411362ndash3
RajaramS Prospects andpromise ofwheatbreedingin the21stcentury Euphytica 20011193-15Rajaram S Role of conventional plant breeding and biotechnology in future wheat
production Turk J Agric Forest 200529105ndash11Rami JF Dufour P Trouche G Fliedel G Mestres C Davrieux F et al Quantitative trait
loci for grain quality productivity morphological and agronomical traits insorghum (Sorghum bicolor L Moench) Theor Appl Genet 199897605ndash16
Rassaa N Salahb HBH Latiri K Thermal responses of durum wheat Triticum durum toearly water stress consequence on leaf and 1047298ower development Plant Biol Pathol2008331(5)363ndash71
Reddy LJ Nigam SN Rao RCN Reddy NS Registration of ICGV 87354 peanut germplasmwith drought tolerance and rust resistance Crop Sci 200141274ndash5
Ribaut JM Ragot M Marker-assisted selection to improve drought adaptation in maize thebackcross approach perspectives limitations and alternatives J Exp Bot 200658351ndash60
Robin S Pathan MS Courtois B La1047297tte R Carandang S Lanceras S et al Mappingosmotic adjustment in an advanced back-cross inbred population of rice Theor
Appl Genet 2003107(7)1288ndash96Romero C Belles JM Vaya JL Serrano R Culianez-Macia FA Expression of the yeast
trehalose-6-phosphate synthase gene in transgenic tobacco plants pleiotropicphenotypes include drought tolerance Planta 1997201293ndash7
Ronde JAD Cress WA Krugerd GHJ Strasserd RJ Van Staden J Photosynthetic responseof transgenic soybean plants containing an Arabidopsis P5CR gene during heat anddrought stress J Plant Physiol 20041611211ndash24
Salvi S Tuberosa R To clone or not to clone plant QTLs present and future challengesTrends Plant Sci 200510297ndash304
Sanchez AC Subudhi PK Rosenow DT Nguyen HT Mapping QTLs associated with droughtresistance in sorghum (Sorghum bicolor L Moench) Plant Mol Biol 200248713ndash26
Saranga Y Menz M Jiang CX Wright RJ Yakir D Paterson AH Genomic dissection of genotypetimesenvironment interactions conferring adaptation of cotton to aridconditions Genome Res 2001111988ndash95
Sari-Gorla M Krajewski P Di Fonzo N Villa M Frova C Genetic analysis of droughttolerance in maize by molecular markers II Plant height and 1047298owering Theor ApplGenet 199999289ndash95
Seki M Narusaka M Ishida J Nanjo T Fujita M Oono Y et al Monitoring the expressionpro1047297les of 7000 Arabidopsis genes under drought cold and high-salinity stressesusing a full-length cDNA microarray Plant J 200231279ndash92
Serraj R Sinclair TR Osmolyte accumulation can it really increase crop yield underdrought conditions Plant Cell Environ 200225333ndash41
Serraj R Krishnamurthy L Kashiwagi J Kumar J Chandra S Crouch JH Variation in roottraits of chickpea (Cicer arietinum L) grown under terminal drought Field CropsRes 200488115ndash27
Serraj R Hash CT Rizvi MHS Sharma A Yadav RS Bidinger FR Recent advances in marker-assisted selection for drought tolerance in pearl millet Plant Prod Sci 20058(3)334ndash7
Shen YG Du BX Zhang WK Zhang JS Chen SY AhCMO regulated by stresses in Atriplexhortensis can improve drought tolerance in transgenic tobacco Theor Appl Genet2002105815ndash21
Shinozaki K Yamaguchi-Shinozaki K Molecular responses to dehydration and lowtemperature differences and cross-talk between two stress signaling pathwaysCurr Opin Plant Biol 20003217ndash23
Singh KB Omar M Saxena MC Johansen C Registration of FLIP 87-59C a drought-tolerant chickpea germplasm line Crop Sci 199636(2)1ndash2
Singh SP Teran H Gutierrez JA Registration of SEA 5 and SEA 13 drought tolerant drybean germplasm Crop Sci 200141276ndash7
Singh SP Teran H Lema M Schwartz HF Miklas PN Registration of white moldresistant dry bean germplasm line A 195 J Plant Registr 2007162ndash3
Sivamani E Bahieldin A Wraith JM Improved biomass productivity and water useef 1047297ciency under water de1047297cit conditions in transgenic wheat constitutivelyexpressing the barley HVA1 gene Plant Sci 20001551ndash9
Skovmand B Reynolds MP DeLacy IH Searching genetic resources for physiologicaltraits with potential for increasing yield In Reynolds MP Ortiz-Monasterio IMcNab A editors Application of Physiology in Wheat Breeding 2001 p 17ndash28
Steele K Novel upland rice variety bred using marker-assisted selection and client-oriented breeding released in Jharkhand India Bangor University 2009
Steele KA Price AH Shashidar HE Witcombe JR Marker-assistedselection to introgressrice QTLs controlling root traitsinto an Indianupland rice variety Theor Appl Genet2006112208ndash21
Steele KA Virk DS Kumar R Prasad SC Witcombe JR Field evaluation of upland ricelines selected for QTLs controlling root traits Field Crops Res 2007101180 ndash6
Sunkar R Kapoor A Zhu JK Post transcriptional induction of two CuZn superoxidedismutase genes in Arabidopsis is mediated by down regulation of miR398 andimportant for oxidative stress tolerance Plant Cell 2006182051ndash65
Talame V Sanguineti MC Chiapparino E Bahri H Ben Salem M Forster BP et alIdenti1047297cation of Hordeum spontaneum QTL alleles improving 1047297eld performance of barley grown under rainfed conditions Ann Appl Bot 2004144309ndash20
Tanner NKCordinO BanroquesJ DoereM Linder PThe Q Motif a newlyidenti1047297ed motif in DEAD box helicases may regulate ATP binding and hydrolysis Mol Cell 200311127ndash38
Taramino G Tarchini R Ferrario S Lee M Pe ME Characterization and mapping of simplesequence repeats (SSRs) in Sorghum bicolor Theor Appl Genet 19979566ndash72
Teulat B Monneveux P Wery J Borriegraves C Souyris I Charrier A et al Relationshipsbetween relative water content andgrowth parameters in barleya QTLstudy NewPhytol 199713799-107
Thi Lang N Chi Buu B Fine mapping for drought tolerance in rice ( Oryza sativa L)Omonrice 2008169-15Till BJ Comai L Henikoff S Tillering and ecotillering for crop improvement In
Varshney RK Tuberosa R editors Genomics-assisted crop improvement genomicsapproaches and platforms vol 1 2007 p 333ndash50
Tondelli A Francia E Barabaschi D Aprile A Skinner JS Stockinger EJ et al Mappingregulatory genes as candidates for cold and drought stress tolerance in barleyTheor Appl Genet 2006112445ndash54
Tuberosa R Salvi S Genomics approaches to improve drought tolerance in cropsTrends Plant Sci 200611405ndash12
Valkoun JJ Wheat pre-breeding using wild progenitors Euphytica 200111917ndash23Vashisht AA Tuteja N Stress responsive DEAD-box helicases a new pathway to
engineer plant stress tolerance J Photochem Photobiol B Biol 200684150 ndash60Vendruscolo ECG Schuster I Pileggi M Scapim CA Molinari HBC Marur CJ et al Stress-
induced synthesis of proline confers tolerance to water de1047297cit in transgenic wheat J Plant Physiol 20071641367ndash76
Vienne D Leonardi A Damerval C Zivy M Genetics of proteome variation for QTL characterization application to drought stress responses in maize J Exp Bot 199950303ndash9
Villareal RL Mujeeb-Kazi A Rajaram S Toro ED Morphological variability in somesynthetic hexaploid wheats derived from Triticum turgidum times T tauschii J GenetBreed 1994487-16
Vinh NT Paterson AH Genome mapping and its implication for stress resistance inplants In Ashraf M Harris PJC editors Abiotic stresses plant resistance throughbreeding and molecular approaches 2005
Vinocur B Altman A Recent advances in engineering plant tolerance to abiotic stressachievements and limitations Curr Opin Biotechnol 200516123ndash32
Wang YJ Hao YJ Zhang ZG Chen T Zhang JS Chen SY Isolation of trehalose-6-phosphate phosphatase gene from tobacco and its functional analysis in yeast cells
J Plant Physiol 2005162215ndash23Wu R Garg A Engineering rice plants with trehalose-producing genes improves
tolerance to drought salt and low temperature ISB News Report 2003Xiao B Huang Y Tang N Xiong L Overexpression of LEA gene in rice improves drought
resistance under 1047297eld conditions Theor Appl Genet 200711535ndash46Xinglai P Sangang X Qiannying P Yinhong S Registration of lsquo Jinmai 50rsquo wheat Crop Sci
200646983ndash5Xiong L Lee H Ishitani M Zhu JK Regulation of osmotic stress-responsive gene
expression by the LOS6ABA1 locus in Arabidopsis J Biol Chem 20022778588ndash96Xu GW Magill CW Shertz KF Hart GE A RFLP linkage map of Sorghum bicolor (L)
Moench Theor Appl Genet 199489139ndash45Xu D Duan X Wang B Hong BHo THD Wu RExpression of a late embryogenesis abundant
protein gene HVA1 from barley confers tolerance to water de1047297cit and salt stress intransgenic rice Plant Physiol 1996110249ndash57
Yadav RS Hash CT Bidinger FR Devos KM Howarth CJ Genomic regions associatedwith grain yield and aspects of post-1047298owering drought tolerance in pearl milletacross stress environments and testers background Euphytica 2004136265ndash77
YamadaM MorishitaH Urano K Shiozaki N Yamaguchi-Shinozaki K ShinozakiK et alEffects of free proline accumulation in petunias under drought stress J Exp Bot2005561975ndash81
Yang WJ Rich PJ Axtell JD Wood KV Bonham CC Ejeta G et al Genotypic variation forglycinebetaine in sorghum Crop Sci 200343162ndash9
Yan-Ying QU Ping MU Xue-Qin L Yu-Xiu T Feng W Hong-Liang Z et al QTL mappingand correlations between leaf water potential and drought resistance in rice underupland and lowland environments Acta Agron Sin 200834(2)198ndash206
Ye XDWu XLZhaoH Frehner MNoumlsberger J Potrykus Iet al Altered fructan accumulationin transgenic Lolium multi 1047298orum plants expressing a Bacillus subtilis sacB gene Plant Cell
Rep 200120205ndash
12Zaharieva M Gaulin E Havaux M Acevedo E Monneveux P Drought and heat responses in
the wild wheat relative Aegilops geniculata Roth potential interest for wheatimprovement Crop Sci 2001411321ndash9
Zhang J Zheng HGAartiA Pantuwan G NguyenTT Tripathy JNet alLocatinggenomicregions associated with components of drought resistance in rice comparativemapping within and across species Theor Appl Genet 200110319ndash29
Zhang GH Su Q An LJ Wu S Characterization and expression of a vacuolar Na+H+
antiporter gene from the monocot halophyte Aeluropus littoralis Plant PhysiolBiochem 200846117ndash26
Zhao J QTLs for oil content and their relationships to other agronomic traits in anEuropeantimesChinese oilseed rape population Germany Diss Grorg-Agust Univer-sity of Goettingen 2002
Zhao J Ren W Zhi D Wang L Xia G Arabidopsis DREB1ACBF3 bestowed transgenic tallfescue increased tolerance to drought stress Plant Cell Rep 2007261521ndash8
Zhou W Li Y Zhao BC Ge RC Shen YZ Wang G Huang ZJ Over-expression of TaSTRG gene improves salt and drought tolerance in rice J Plant Physiol 200915(166)1660ndash70
and cost-effective Molecular mapping and analysis of QTL have been
carried out for a number of qualitative and quantitative traits including
stress tolerance which has undoubtedly resulted in a great magnitude
of knowledge and better understanding of the causal genetic phenom-
ena that regulate these traits However limited success has resulted by
using this knowledge to manipulate genes in an effective way for the
improvement of a speci1047297c trait in crops although some crop cultivars
developed through using these tools have performed extremely well
under1047297
eld stressconditions However there are a variety of reasons forthe limited success in terms of achieving enhanced drought tolerance
using the marker-assisted breeding approach For example a major
dif 1047297culty confronting the scientists is the challenge of precise QTL
identi1047297cation In addition a substantial genetictimes environment interac-
tion inconsistent repeatability large number of genes regulating yield
and invalid use of mapping populations have hampered the pursuits
involving mapping of QTL for enhanced drought stress tolerance
Despite these other factors also impede the application of QTL for
genetic improvement of a trait For example due to unfavorable
epistatic interaction it is hard to transfer the effects of a desired allele to
an elite background material (Podlich et al 2004 Collins et al 2008)
Furthermore in some cases QTL from a speci1047297c background do not
show signi1047297cant effects or cease completely in different backgrounds
even under analogous growth conditions (Cho and Hong 2006 Collins
et al 2008) This makes the utilization of QTL more intricate
Transformation of the knowledge acquired from QTL-oriented molec-
ular studies into a well-de1047297ned upshot for the stakeholders is one of the
key challenges confronting the breeders Certainly a multidisciplinary
approach including more speci1047297cally the identi1047297cation of QTL mediated
signal transduction in response to stresses needs to be adopted
Furthermore although QTL cloning procedures are unwieldy they can
lead to a meaningful outcome because a cloned QTL can offer a reliable
markerfor MABas well as furnish an outlinefor thedetection of superior
allelic variants in crop species (Till et al 2007)
Genetic engineering (Transgenic approach) offers a promise whereby
one can expect a substantial improvement in a desired trait within the
shortest stretch of time Engineering genes encoding organic osmolytes
plant growth regulators antioxidants late embryogenesis abundant pro-
teins and transcription factors has resulted into transgenic lines whichhave out-performed under controlled stress conditions In most cases the
claims made by the researchers in terms of performance of transgenic
lines tested under controlled conditions are considerably in1047298ated
However with the exception of a very few notable cases most of the
transgenic lines of differentcrops have been rarely1047297eld-tested Thushow
far these transgenic lines perform under 1047297eld stress conditions is not
known because under natural1047297eld conditions a linecultivar has to face a
multitude of environmental factors other than the drought stress Thus
appraisal of performance of a transgenic line under natural 1047297eld con-
ditionsis mandatory forthe stakeholdersperspective It is also imperative
to note that most of the drought tolerant transgenic lines have been
developed using a single gene transformation which may not be as
productive as if it had been developed using transformation of many
genes Thus transferring a number of prominent genes effectively in-volved in stress tolerance to transgenic plants seems to be a logical
approach Although a large number of genes appear to be involved in
stress tolerance and most of them have been fully characterized the
function of many of them in the mechanism of stress tolerance is yet to be
investigated
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Ashraf M Breeding for salinity tolerance in plants Crit Rev Plant Sci 19941317 ndash42Ashraf M Akram NA Improving salinity tolerance of plants through conventional
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Ashraf M Foolad MR Roles of glycinebetaine and proline in improving plant abioticstress resistance Environ Exp Bot 200759206ndash16
Ashraf M Athar HR Harris PJC Kwon TR Some prospective strategies for improvingcrop salt tolerance Adv Agron 20089745-110
Asins MJ Present and future of quantitative trait locus analysis in plant breeding PlantBreed 2002121281ndash91
Babu RC Nguyen BD Chamarerk V Shanmugasundaram P Chezhian P Jeyaprakash P et alGeneticanalysis of droughtresistance in riceby molecular markers association betweensecondary traits and 1047297eld performance Crop Sci 2003431457ndash69
Babu RC Zhang J Blum A Ho THD Wu R Nguyen HT HVA1 a LEA gene from barleyconfers dehydration tolerance in transgenic rice( Oryzasativa L)via cell membraneprotection Plant Sci 2004166855ndash62
Badawi GH Kawano N Yamauchi Y Over-expression of ascorbate peroxidase in
tobacco chloroplasts enhances the tolerance to salt stress and water de1047297cit PhysiolPlant 2004121231ndash8Badu-Apraku B Yallou CG Registration of striga-resistant and drought tolerant tropical
early maize populations TZE-W Pop DT STR C4 and TZE-Y Pop DT STR C4 J PlantRegistr 20093(1)86ndash90
Badu-Apraku B Menkir A Kling JG Fakorede MAB Registration of 16 striga resistantearly maturing tropical maize inbred lines Crop Sci 2006461410ndash1
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Baumlnziger M Setimela PS Hodson D Vivek B Breeding for improved drought tolerance inmaize adapted to southern Africa Proceedings of the 4th International Crop ScienceCongress Brisbane Australia Published on CDROM 2004 26 Sep ndash 1 Oct
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traits in the Mediterranean environment identi1047297ed in recombinant inbred lines of the cross lsquoArtarsquo H spontaneum 41-1 Theor Appl Genet 20031071215ndash25
Bergman JW Riveland NR Flynn CR Carlson GR Wichman DM Registration of lsquoMorlinrsquo
saf 1047298ower Crop Sci 2001411640Bernier J Kumar A Venuprasad R Spaner D Atlin G A large-effect QTL for
grain yield under reproductive-stage drought stress in upland rice Crop Sci200747507ndash18
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Bernier J Serraj R Kumar A Venuprasad R Impa S Gowdaa RPV et al The large-effectdrought-resistance QTL qtl121 increases water uptake in upland rice Field CropsRes 200911039ndash46
Bhatnagar-Mathur P ReddyDS Lavanya M Yamaguchi-Shinozaki K Sharma KK Stress-inducible expression of Arabidopsis thaliana DREB1A in transgenic peanut ( Arachishypogaea L) increases transpiration ef 1047297ciency under water-limiting conditionsPlant Cell Rep 2007262071ndash82
Bhatnagar-Mathur P Devi MJ Vadez V Sharma KK Differential antioxidative responses intransgenic peanut bear no relationship to their superior transpiration ef 1047297ciency underdrought stress J Plant Physiol 2009166(11)1207ndash17 doi101016jjplph200901001
Bidinger FR Serraj R Rizvi SMH Howarth C Yadav RS Hash CT Field evaluation of droughttolerance QTL effects on phenotype and adaptation in pearl millet [ Pennisetum glaucum(L) R Br] topcross hybrids Field Crops Res 200594(1)14ndash32
Bidinger FR Nepolean T Hash CT Yadav RS Howarth CJ Identi1047297cation of QTLs for grainyield of pearl millet (Pennisetum glaucum (L) R Br) in environments with variablemoisture during grain 1047297lling Crop Sci 200747969ndash80
Bowers JE Abbey C Anderson S Chang C Draye X A high-density geneticrecombination map of sequence-tagged sites for Sorghum as a framework forcomparative structural and evolutionary genomics of tropical grains and grassesGenetics 2003165367ndash86
Brick MA Ogg JB Singh SP Schwartz HF Johnson JJ Pastor-Corrales MA Registration of drought-tolerant rust-resistant high-yielding pinto bean germplasm lineCO46348 J Plant Registr 20082(2)120ndash4
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Busk PK Pages M Regulation of abscisic acid induced transcription Plant Mol Biol199837425ndash35
Carena MJ Wanner DW Development of genetically broad-based inbred lines of maizefor early-maturing (70-80RM) hybrids J Plant Registr 20093107ndash11
Cash SDBrucknerPL Wichman DMKephart KD Berg JEBoynerR et alRegistration of Willow Creek forage wheat J Plant Registr 20093(2)185ndash90
Cattivelli L Rizza F Badeck FW Mazzucotelli E Francia AMEM Mare AT et al Droughttolerance improvement in crop plants an integrated view from breeding to genomics
Field Crops Res 20081051-14Chen M Wang QY Cheng XG Xu ZS Li LC Ye XG et al GmDREB2 a soybean DRE-
binding transcription factor conferred drought and high-salt tolerance intransgenic plants Biochem Biophys Res Commun 2007353299ndash305
Cheng Z Targolli J Huang X Wu R Wheat LEA genes PMA80 and PMA1959 enhancedehydration tolerance of transgenic rice (Oryza sativa L) Mol Breed 20021071ndash82
ChoEK Hong ChB Over-expression of tobacco NtHSP70-1 contributes to drought-stresstolerance in plants Plant Cell Reports 200625349ndash58
Close TJ Dehydrins a commonality in the response of plants to dehydration and lowtemperature Physiol Plant 1997100291ndash6
Collins NC Tardieu F Tuberosa R Quantitative trait loci and crop performance underabiotic stress where do we stand Plant Physiol 2008147469ndash86
Concept Note Combining breeding and biotechnology to develop water ef 1047297cient maizefor Africa (WEMA) Afr Agric Technol Foundation
Courtois BShen L Petalcorin WCarandang S MauleonR LiZ Locating QTLs controllingconstitutive root traits in the rice population IAC 165-Co39 Euphytica 2003134335ndash45
Da-hong L Hui L Yang YL Ping-ping Z Jian-sheng L Down-regulated expression of RACK1geneby RNAinterference enhancesdroughttolerancein rice RiceSci 200916(1)14ndash20
Dalal M Tayal D Chinnusamy V Bansala KC Abiotic stress and ABA-inducible group 4 LEAfrom Brassicanapus plays a key rolein saltand droughttolerance J Biotechnol 2009139137ndash45
Ding Z Li S An X Liu X Qin H Wang D Transgenic expression of MYB15 confers enhancedsensitivity to abscisic acid and improved drought tolerance in Arabidopsis thaliana
J Genet Genomics 20093617ndash29Dubouzet JG Sakuma Y Ito Y Kasuga M Dubouzet EG Miura S et al OsDREB genes in
rice Oryza sativa L encode transcription activators that function in drought high-salt- and cold-responsive gene expression Plant J 200333751ndash63
El-Far IA Allan AY Responses of some wheat cultivars to sowing methods and droughtat different stages of growth Assuit J Agric Sci 199526(1)267ndash77
Eltayeb AE Kawano N Badawi GH Kaminaka H Sanekata T Shibahara T et alOverexpression of monodehydroascorbate reductase in transgenic tobacco confersenhanced tolerance to ozonesalt and polyethyleneglycol stresses Planta 2007225(5)1255ndash64
Falconer DS Introduction to quantitative genetics London New York Longman 1989Feng-ling FU Zhi-Lei F Shi-bing G Shu-feng Z Wan-chen L Evaluation and quantitative
inheritanceof several drought-relativetraits in maize Agric SciChina 20087(3)280ndash90Finkelstein R Gampala S Rock C Abscisic acid signaling in seeds and seedlings Plant
Cell 20021415ndash45Flowers TJ Improving crop salt tolerance J Exp Bot 200455307ndash19Fujita Y Fujita M Satoh R Maruyama K Parvez MM Seki M et al AREB1 is a transcription
Giraudat J Parcy F Bertauche N Gosti F Leung J Morris PC et al Current advances inabscisic acid action and signalling Plant Mol Biol 1994261557ndash77
Gorbalenya AE Koonin EV Helicases amino acid sequence comparisons and structurendashfunction relationships Curr Opin Struct Biol 19933419ndash29
Gosal SS Wani SH Kang MS Biotechnology and drought tolerance J Crop Improvement20092319ndash54
Gubis J Vaňkovaacute R Červenaacute V Draguacuteňovaacute M Hudcovicovaacute M Lichtnerovaacute H et alTransformedtobacco plants with increased toleranceto droughtSouthAfr J Bot200773505ndash11
GuoO Zhang J GaoQ Xing SLi F Wang WDrought tolerancethrough over-expressionof mono ubiquitin in transgenic tobacco J Plant Physiol 20081651745 ndash55
Haley SD Johnson JJ Peairs FB Quick JS Stromberger JA Clayshulte SR et al Registration of lsquoRipperrsquo wheat J Plant Registr 200711ndash6
Han SE Park SR Kwon HB Yi BY Lee GB Byun MO Genetic engineering of drought-resistant tobacco plants by introducingthe trehalose phosphorylase (TP) gene fromPleurotus sajor-caju Plant Cell Tissue Organ Cult 200582151ndash8
Harris K Klein R Mullet J Sorghum stay-green QTL individually reduces post-1047298oweringdrought-induced leaf senescence J Exp Bot 200758327ndash38
Hong BS Zong-Suo L Ming-An S LEA proteins in higher plants structure functiongene expression and regulation Colloids Surf B Biointerf 200545131ndash5
Howarth CJ Yadav RS Successful marker assisted selection for drought tolerance anddisease resistance in pearl milletIGER Innovations 2002
Humphreys MO Humphreys MW Breeding for stress resistance general principles InAshraf M Harris PJC editors Abiotic stresses plant resistance through breedingand molecular approaches 2005 p 19ndash46
Ingram J Bartels D The molecular basis of dehydration tolerance in plants Ann RevPlant Physiol Plant Mol Biol 199647377ndash403
Jaglo-Ottosen KR Gilmour SJ Zarka DG Schabenberger O Thomashow MF ArabidopsisCBF1 overexpression induces COR genes and enhances freezing tolerance Science1998280104ndash6
Jang IC Oh SJ Seo JS Choi WB Song SI Kim CH et al Expression of a bifunctional fusion of the Escherichia coli genes for trehalose-6-phosphate synthase and trehalose-6-phosphate phosphatase in transgenic rice plants increases trehalose accumulationandabiotic stresstolerancewithoutstuntinggrowth Plant Physiol2003131516ndash24
Jeanneau M Gerentes D Foueillassar X Zivy M Vidal J Toppan A et al Improvement of drought tolerance in maize towards the functional validation of the Zm-Asr1 gene andincrease of water use ef 1047297ciency by over-expressing C4-PEPC Biochimie 2002841127ndash35
Jenson KB Palazzo AJ Waldron BL Bushman BS Registration of lsquoFirstStrikersquo Slenderwheatgrass J Plant Registr 2007124ndash5
Joung-youn K Hyung-in C Min-young I Soo-young K Arabidopsis basic leucine zipperproteins that mediate stress-responsive abscisic acid signaling Plant Cell 20024343ndash57
Juskiw PE Helm JH Oro M Nyachiro JM Salmon DF Registration of lsquoBentleyrsquo barley
J Plant Registr 20093(2)119ndash23Kalamaki MS Alexandrou D Lazari D Merkouropoulos G Fotopoulos V Pateraki I et al
Over-expression of a tomato N-acetyl-L -glutamate synthase gene (SlNAGS1) in Arabidopsis thaliana results in high ornithine levels and increased tolerance in saltand drought stresses J Exp Bot 200960(6)1859ndash71
Kamoshita A Babu CR Boopathi NM Fukai S Phenotypic and genotypic analysis of drought-resistance traits for development of rice cultivars adapted to rainfedenvironments Field Crops Res 2008109(103)1-23 doi101016jfcr200806010
KarakasB Ozias-AkinsP Stushnoff C SuefferheldM Rieger M Salinityand drought toleranceof mannitol-accumulating transgenic tobacco Plant Cell Environ 199720609ndash16
Karim S Aronsson H Ericson H Pirhonen M Leyman B Welin B et al Improved droughttolerance without undesired side effects in transgenic plants producing trehalose PlantMol Biol 200764371ndash86
Kasuga M Liu Q Miura S Yamaguchi-Shinozaki K Shinozaki K Improving plant droughtsaltand freezing tolerance by gene transfer of a single stress-inducible transcription factorNat Biotechnol 199917287ndash91
Kindiger M Gaub H Hasegawac M Katsurab Y Ueyamad K Gotob S et al Registrationof lsquoNanryorsquo tall fescue Crop Sci 2006461815ndash6
Kong I Dong J HartGE Characteristics linkage mappositionsand allelic differentiationof Sorghum bicolour (L) Moench by DNA simple-sequence repeats (SSRs) TheorAppl Genet 2000101438ndash48
Kumar R Venuprasad R Atlin GN Genetic analysis of rainfed lowland rice droughttolerance under naturally-occurring stress in eastern India heritability and QTL effects Field Crops Res 200710342ndash52
La1047297tte HR Price AH Courtois B Yield response to water de1047297cit in an upland ricemapping population associations among traits and genetic markers Theor ApplGenet 20041091237ndash46
Lal S Gulyani V Khurana P Overexpressionof HVA1 gene frombarley generates tolerance tosalinity and water stress in transgenic mulberry (Morus indica) Transgenic Res 200817
(4)651ndash
63Lanceras J Pantuwan G Jongdee B Toojinda T Quantitative trait loci associated withdrought tolerance at reproductive stage in rice Plant Physiol 2004135384ndash99
Levi A Ovnat L Paterson AH Saranga Y Photosynthesis of cotton near-isogenic linesintrogressed with QTLs for productivity and drought related traits Plant Sci 2009a17788ndash96
Levi A Paterson AH Barak V Yakir D Wang B Chee PW et al Field evaluation of cottonnear-isogenic lines introgressed with QTLs for productivity and drought relatedtraits Mol Breed 2009b23179ndash95
Liu Q Kasuga M Sakuma Y Abe H Miura S Yamaguchi- Shinozaki K et al Twotranscription factors DREB1 and DREB2 with an EREBPAP2 DNA binding domainseparate two cellular signal transduction pathways in drought-and low-temper-ature-responsive gene expression respectively in Arabidopsis Plant Cell 1998101391ndash406
Liu X Hua X Guo J Qi D Wang L Liu Z et al Enhanced tolerance to drought stress intransgenic tobacco plants overexpressing Biotechnol Lett 2008301275ndash80
Liu X Wanga Z Wanga L Wua R Phillips J Deng X LEA 4 group genes from theresurrection plant Boea hygrometrica confer dehydration tolerance in transgenictobacco Plant Sci 200917690ndash8
Luchi S Kobayashi M Taji T Naramoto M Seki M Kato T et al Regulation of droughttolerance by gene manipulation of 9-cis-epoxycarotenoid dioxygenase a keyenzyme in abscisic acid biosynthesis in Arabidopsis Plant J 200127325ndash33
Luo Y Liu YB Dong YX Gao XQ Zhang YS Expression of a putative alfalfa helicase increasestolerance to abioticstress in Arabidopsis by enhancing the capacities for ROS scavengingand osmotic adjustment J Plant Physiol 2009166385ndash94
MacLean JLDaweDC Hardy B Hettel GPRiceAlmanac sourcebook for themostimportanteconomic activity on Earth3rd ed Wallingford England CABI Publishing 2002
Mansour MMF Nitrogen containing compounds and adaptation of plants to salinitystress Biol Plant 200043491ndash500
McKersie BD Bowley SR Harjanto E Leprince O Water-de1047297cit tolerance and 1047297eldperformance of transgenic alfalfa overexpressing superoxide dismutase Plant Physiol19961111177ndash81
McKersie BD Murnaghan J Bowley SR Manipulating freezing tolerance in transgenicplants Acta Physiol Plant 199719485ndash95
Miranda JA Avonce N Suaacuterez R Thevelein JM Dijck PV Iturriaga G A bifunctional TPS-TPP enzyme from yeast confers tolerance to multiple and extreme abiotic-stressconditions in transgenic Arabidopsis Planta 2007226(6)1411ndash21
Mohammadi M Taleei A Zeinali H Naghavi MR Ceccarelli S Grando Baum M QTL analysis for phenologic traits in doubled haploid population of barley Int J AgricBiol 20057(5)820ndash3
Mohanty A Kathuria H Ferjani A Sakamoto A Mohanty P Murata N et al Transgenicsof an elite indica rice variety Pusa Basmati 1 harbouring the codA gene are highlytolerant to salt stress Theor Appl Genet 200210651ndash7
Noaman MM El Sayad AA Asaad FA El Sherbini AM El Bawab AO El Moselhi MA et alRegistration of lsquoGiza 126rsquo barley Crop Sci 199535(6)1710
Noaman MM Ahmed IA El-Sayed AA Abo-El-Enin RA El-Gamal AS El-Sherbiny AM et alRegistration of lsquoGiza 2000rsquo drought-tolerant six-rowed barley for rainfed and newreclaimed areas in Egypt Crop Sci 200747440
Obert DE Evans CP Wesenberg DM Windes JM Erickson CA Jackson EW et alRegistration of lsquoLenetahrsquo spring barley J Plant Registr 20082(2)85ndash7
Oh SJ Jeong JS Kim EH Yi NR Yi SI Jang IC et al Matrix attachment region fromthe chicken lysozyme locus reduces variability in transgene expression andconfers copy number-dependence in transgenic rice plants Plant Cell Rep20054145ndash54
Oh SJ Kim YS Kwon C Park HK Jeong JS Kim JK Overexpression of the transcriptionfactor AP37 in rice improves grain yield under drought conditions Plant Physiol
20092191ndash
200 doi101104pp109137554Owttrim GW RNA helicases and abiotic stress Nucleic AcidsRes 200634(11)3220ndash30Park BJ Liu Z Kanno A Kameya T Increased tolerance to salt and water de1047297cit stress in
transgenic lettuce (Lactuca sativa L) by constitutive expression of LEA PlantGrowth Regul 2005a45165ndash71
Park BJ Liu Z Kanno A Kameya T Genetic improvement of Chinese cabbage for salt anddroughttolerance by constitutiveexpression of a B napus LEAgenePlantSci 2005b169553ndash8
Pastori GM Foyer CH Common componentsnetworks and pathways of crosstoleranceto stress The central role of ldquoredoxrdquo and abscisic acid-mediated controls PlantPhysiol 2002129460ndash8
PellegrineschiA ReynoldsM PachecoM Brito RMAlmeraya R Yamaguchi-ShinozakiK et alStress-induced expression in wheat of the Arabidopsis thaliana DREB1Agenedelayswaterstress symptoms under greenhouse conditions Genome 200447493ndash500
Perl A Perl-Treves R Galili S Aviv D Shalgi E Malkin S et al Enhanced oxidative-stressdefense in transgenic potato overexpressing tomato Cu Zn superoxide dismutaseTheor Appl Genet 199385568ndash76
Podlich DW Winkler CR Cooper M Mapping as you go an effective approach formarker-assisted selection of complex traits Crop Sci 2004441560ndash71
182 M Ashraf Biotechnology Advances 28 (2010) 169ndash183
Pratt RC Casey MA Registration of maize germplasm line Oh605 Crop Sci 2006461004ndash5Qin F Kakimoto M Sakuma Y Maruyama K Osakabe Y Tran LSP et al Regulation and
functional analysis of ZmDREB2A in response to drought and heat stress in Zea mays LPlant J 20075054ndash69
Quan R Shang M Zhang H Zhao Y Zhang J Engineering of enhanced glycinebetainesynthesis improves drought tolerance in maize Plant Biotechnol J 20042477ndash86
Quarrie SA Gulli M Calestani C Steed A Marmiroli N Location of a gene regulatingdrought-induced abscisic acid production on the long arm of chromosome 5A of wheat Theor Appl Genet 199489794ndash800
QuickJS Stromberger JA Clayshulte S Clifford B Johnson JJ Peairs FB et al Registrationof lsquoPrairie Redrsquo wheat Crop Sci 2001411362ndash3
RajaramS Prospects andpromise ofwheatbreedingin the21stcentury Euphytica 20011193-15Rajaram S Role of conventional plant breeding and biotechnology in future wheat
production Turk J Agric Forest 200529105ndash11Rami JF Dufour P Trouche G Fliedel G Mestres C Davrieux F et al Quantitative trait
loci for grain quality productivity morphological and agronomical traits insorghum (Sorghum bicolor L Moench) Theor Appl Genet 199897605ndash16
Rassaa N Salahb HBH Latiri K Thermal responses of durum wheat Triticum durum toearly water stress consequence on leaf and 1047298ower development Plant Biol Pathol2008331(5)363ndash71
Reddy LJ Nigam SN Rao RCN Reddy NS Registration of ICGV 87354 peanut germplasmwith drought tolerance and rust resistance Crop Sci 200141274ndash5
Ribaut JM Ragot M Marker-assisted selection to improve drought adaptation in maize thebackcross approach perspectives limitations and alternatives J Exp Bot 200658351ndash60
Robin S Pathan MS Courtois B La1047297tte R Carandang S Lanceras S et al Mappingosmotic adjustment in an advanced back-cross inbred population of rice Theor
Appl Genet 2003107(7)1288ndash96Romero C Belles JM Vaya JL Serrano R Culianez-Macia FA Expression of the yeast
trehalose-6-phosphate synthase gene in transgenic tobacco plants pleiotropicphenotypes include drought tolerance Planta 1997201293ndash7
Ronde JAD Cress WA Krugerd GHJ Strasserd RJ Van Staden J Photosynthetic responseof transgenic soybean plants containing an Arabidopsis P5CR gene during heat anddrought stress J Plant Physiol 20041611211ndash24
Salvi S Tuberosa R To clone or not to clone plant QTLs present and future challengesTrends Plant Sci 200510297ndash304
Sanchez AC Subudhi PK Rosenow DT Nguyen HT Mapping QTLs associated with droughtresistance in sorghum (Sorghum bicolor L Moench) Plant Mol Biol 200248713ndash26
Saranga Y Menz M Jiang CX Wright RJ Yakir D Paterson AH Genomic dissection of genotypetimesenvironment interactions conferring adaptation of cotton to aridconditions Genome Res 2001111988ndash95
Sari-Gorla M Krajewski P Di Fonzo N Villa M Frova C Genetic analysis of droughttolerance in maize by molecular markers II Plant height and 1047298owering Theor ApplGenet 199999289ndash95
Seki M Narusaka M Ishida J Nanjo T Fujita M Oono Y et al Monitoring the expressionpro1047297les of 7000 Arabidopsis genes under drought cold and high-salinity stressesusing a full-length cDNA microarray Plant J 200231279ndash92
Serraj R Sinclair TR Osmolyte accumulation can it really increase crop yield underdrought conditions Plant Cell Environ 200225333ndash41
Serraj R Krishnamurthy L Kashiwagi J Kumar J Chandra S Crouch JH Variation in roottraits of chickpea (Cicer arietinum L) grown under terminal drought Field CropsRes 200488115ndash27
Serraj R Hash CT Rizvi MHS Sharma A Yadav RS Bidinger FR Recent advances in marker-assisted selection for drought tolerance in pearl millet Plant Prod Sci 20058(3)334ndash7
Shen YG Du BX Zhang WK Zhang JS Chen SY AhCMO regulated by stresses in Atriplexhortensis can improve drought tolerance in transgenic tobacco Theor Appl Genet2002105815ndash21
Shinozaki K Yamaguchi-Shinozaki K Molecular responses to dehydration and lowtemperature differences and cross-talk between two stress signaling pathwaysCurr Opin Plant Biol 20003217ndash23
Singh KB Omar M Saxena MC Johansen C Registration of FLIP 87-59C a drought-tolerant chickpea germplasm line Crop Sci 199636(2)1ndash2
Singh SP Teran H Gutierrez JA Registration of SEA 5 and SEA 13 drought tolerant drybean germplasm Crop Sci 200141276ndash7
Singh SP Teran H Lema M Schwartz HF Miklas PN Registration of white moldresistant dry bean germplasm line A 195 J Plant Registr 2007162ndash3
Sivamani E Bahieldin A Wraith JM Improved biomass productivity and water useef 1047297ciency under water de1047297cit conditions in transgenic wheat constitutivelyexpressing the barley HVA1 gene Plant Sci 20001551ndash9
Skovmand B Reynolds MP DeLacy IH Searching genetic resources for physiologicaltraits with potential for increasing yield In Reynolds MP Ortiz-Monasterio IMcNab A editors Application of Physiology in Wheat Breeding 2001 p 17ndash28
Steele K Novel upland rice variety bred using marker-assisted selection and client-oriented breeding released in Jharkhand India Bangor University 2009
Steele KA Price AH Shashidar HE Witcombe JR Marker-assistedselection to introgressrice QTLs controlling root traitsinto an Indianupland rice variety Theor Appl Genet2006112208ndash21
Steele KA Virk DS Kumar R Prasad SC Witcombe JR Field evaluation of upland ricelines selected for QTLs controlling root traits Field Crops Res 2007101180 ndash6
Sunkar R Kapoor A Zhu JK Post transcriptional induction of two CuZn superoxidedismutase genes in Arabidopsis is mediated by down regulation of miR398 andimportant for oxidative stress tolerance Plant Cell 2006182051ndash65
Talame V Sanguineti MC Chiapparino E Bahri H Ben Salem M Forster BP et alIdenti1047297cation of Hordeum spontaneum QTL alleles improving 1047297eld performance of barley grown under rainfed conditions Ann Appl Bot 2004144309ndash20
Tanner NKCordinO BanroquesJ DoereM Linder PThe Q Motif a newlyidenti1047297ed motif in DEAD box helicases may regulate ATP binding and hydrolysis Mol Cell 200311127ndash38
Taramino G Tarchini R Ferrario S Lee M Pe ME Characterization and mapping of simplesequence repeats (SSRs) in Sorghum bicolor Theor Appl Genet 19979566ndash72
Teulat B Monneveux P Wery J Borriegraves C Souyris I Charrier A et al Relationshipsbetween relative water content andgrowth parameters in barleya QTLstudy NewPhytol 199713799-107
Thi Lang N Chi Buu B Fine mapping for drought tolerance in rice ( Oryza sativa L)Omonrice 2008169-15Till BJ Comai L Henikoff S Tillering and ecotillering for crop improvement In
Varshney RK Tuberosa R editors Genomics-assisted crop improvement genomicsapproaches and platforms vol 1 2007 p 333ndash50
Tondelli A Francia E Barabaschi D Aprile A Skinner JS Stockinger EJ et al Mappingregulatory genes as candidates for cold and drought stress tolerance in barleyTheor Appl Genet 2006112445ndash54
Tuberosa R Salvi S Genomics approaches to improve drought tolerance in cropsTrends Plant Sci 200611405ndash12
Valkoun JJ Wheat pre-breeding using wild progenitors Euphytica 200111917ndash23Vashisht AA Tuteja N Stress responsive DEAD-box helicases a new pathway to
engineer plant stress tolerance J Photochem Photobiol B Biol 200684150 ndash60Vendruscolo ECG Schuster I Pileggi M Scapim CA Molinari HBC Marur CJ et al Stress-
induced synthesis of proline confers tolerance to water de1047297cit in transgenic wheat J Plant Physiol 20071641367ndash76
Vienne D Leonardi A Damerval C Zivy M Genetics of proteome variation for QTL characterization application to drought stress responses in maize J Exp Bot 199950303ndash9
Villareal RL Mujeeb-Kazi A Rajaram S Toro ED Morphological variability in somesynthetic hexaploid wheats derived from Triticum turgidum times T tauschii J GenetBreed 1994487-16
Vinh NT Paterson AH Genome mapping and its implication for stress resistance inplants In Ashraf M Harris PJC editors Abiotic stresses plant resistance throughbreeding and molecular approaches 2005
Vinocur B Altman A Recent advances in engineering plant tolerance to abiotic stressachievements and limitations Curr Opin Biotechnol 200516123ndash32
Wang YJ Hao YJ Zhang ZG Chen T Zhang JS Chen SY Isolation of trehalose-6-phosphate phosphatase gene from tobacco and its functional analysis in yeast cells
J Plant Physiol 2005162215ndash23Wu R Garg A Engineering rice plants with trehalose-producing genes improves
tolerance to drought salt and low temperature ISB News Report 2003Xiao B Huang Y Tang N Xiong L Overexpression of LEA gene in rice improves drought
resistance under 1047297eld conditions Theor Appl Genet 200711535ndash46Xinglai P Sangang X Qiannying P Yinhong S Registration of lsquo Jinmai 50rsquo wheat Crop Sci
200646983ndash5Xiong L Lee H Ishitani M Zhu JK Regulation of osmotic stress-responsive gene
expression by the LOS6ABA1 locus in Arabidopsis J Biol Chem 20022778588ndash96Xu GW Magill CW Shertz KF Hart GE A RFLP linkage map of Sorghum bicolor (L)
Moench Theor Appl Genet 199489139ndash45Xu D Duan X Wang B Hong BHo THD Wu RExpression of a late embryogenesis abundant
protein gene HVA1 from barley confers tolerance to water de1047297cit and salt stress intransgenic rice Plant Physiol 1996110249ndash57
Yadav RS Hash CT Bidinger FR Devos KM Howarth CJ Genomic regions associatedwith grain yield and aspects of post-1047298owering drought tolerance in pearl milletacross stress environments and testers background Euphytica 2004136265ndash77
YamadaM MorishitaH Urano K Shiozaki N Yamaguchi-Shinozaki K ShinozakiK et alEffects of free proline accumulation in petunias under drought stress J Exp Bot2005561975ndash81
Yang WJ Rich PJ Axtell JD Wood KV Bonham CC Ejeta G et al Genotypic variation forglycinebetaine in sorghum Crop Sci 200343162ndash9
Yan-Ying QU Ping MU Xue-Qin L Yu-Xiu T Feng W Hong-Liang Z et al QTL mappingand correlations between leaf water potential and drought resistance in rice underupland and lowland environments Acta Agron Sin 200834(2)198ndash206
Ye XDWu XLZhaoH Frehner MNoumlsberger J Potrykus Iet al Altered fructan accumulationin transgenic Lolium multi 1047298orum plants expressing a Bacillus subtilis sacB gene Plant Cell
Rep 200120205ndash
12Zaharieva M Gaulin E Havaux M Acevedo E Monneveux P Drought and heat responses in
the wild wheat relative Aegilops geniculata Roth potential interest for wheatimprovement Crop Sci 2001411321ndash9
Zhang J Zheng HGAartiA Pantuwan G NguyenTT Tripathy JNet alLocatinggenomicregions associated with components of drought resistance in rice comparativemapping within and across species Theor Appl Genet 200110319ndash29
Zhang GH Su Q An LJ Wu S Characterization and expression of a vacuolar Na+H+
antiporter gene from the monocot halophyte Aeluropus littoralis Plant PhysiolBiochem 200846117ndash26
Zhao J QTLs for oil content and their relationships to other agronomic traits in anEuropeantimesChinese oilseed rape population Germany Diss Grorg-Agust Univer-sity of Goettingen 2002
Zhao J Ren W Zhi D Wang L Xia G Arabidopsis DREB1ACBF3 bestowed transgenic tallfescue increased tolerance to drought stress Plant Cell Rep 2007261521ndash8
Zhou W Li Y Zhao BC Ge RC Shen YZ Wang G Huang ZJ Over-expression of TaSTRG gene improves salt and drought tolerance in rice J Plant Physiol 200915(166)1660ndash70
and cost-effective Molecular mapping and analysis of QTL have been
carried out for a number of qualitative and quantitative traits including
stress tolerance which has undoubtedly resulted in a great magnitude
of knowledge and better understanding of the causal genetic phenom-
ena that regulate these traits However limited success has resulted by
using this knowledge to manipulate genes in an effective way for the
improvement of a speci1047297c trait in crops although some crop cultivars
developed through using these tools have performed extremely well
under1047297
eld stressconditions However there are a variety of reasons forthe limited success in terms of achieving enhanced drought tolerance
using the marker-assisted breeding approach For example a major
dif 1047297culty confronting the scientists is the challenge of precise QTL
identi1047297cation In addition a substantial genetictimes environment interac-
tion inconsistent repeatability large number of genes regulating yield
and invalid use of mapping populations have hampered the pursuits
involving mapping of QTL for enhanced drought stress tolerance
Despite these other factors also impede the application of QTL for
genetic improvement of a trait For example due to unfavorable
epistatic interaction it is hard to transfer the effects of a desired allele to
an elite background material (Podlich et al 2004 Collins et al 2008)
Furthermore in some cases QTL from a speci1047297c background do not
show signi1047297cant effects or cease completely in different backgrounds
even under analogous growth conditions (Cho and Hong 2006 Collins
et al 2008) This makes the utilization of QTL more intricate
Transformation of the knowledge acquired from QTL-oriented molec-
ular studies into a well-de1047297ned upshot for the stakeholders is one of the
key challenges confronting the breeders Certainly a multidisciplinary
approach including more speci1047297cally the identi1047297cation of QTL mediated
signal transduction in response to stresses needs to be adopted
Furthermore although QTL cloning procedures are unwieldy they can
lead to a meaningful outcome because a cloned QTL can offer a reliable
markerfor MABas well as furnish an outlinefor thedetection of superior
allelic variants in crop species (Till et al 2007)
Genetic engineering (Transgenic approach) offers a promise whereby
one can expect a substantial improvement in a desired trait within the
shortest stretch of time Engineering genes encoding organic osmolytes
plant growth regulators antioxidants late embryogenesis abundant pro-
teins and transcription factors has resulted into transgenic lines whichhave out-performed under controlled stress conditions In most cases the
claims made by the researchers in terms of performance of transgenic
lines tested under controlled conditions are considerably in1047298ated
However with the exception of a very few notable cases most of the
transgenic lines of differentcrops have been rarely1047297eld-tested Thushow
far these transgenic lines perform under 1047297eld stress conditions is not
known because under natural1047297eld conditions a linecultivar has to face a
multitude of environmental factors other than the drought stress Thus
appraisal of performance of a transgenic line under natural 1047297eld con-
ditionsis mandatory forthe stakeholdersperspective It is also imperative
to note that most of the drought tolerant transgenic lines have been
developed using a single gene transformation which may not be as
productive as if it had been developed using transformation of many
genes Thus transferring a number of prominent genes effectively in-volved in stress tolerance to transgenic plants seems to be a logical
approach Although a large number of genes appear to be involved in
stress tolerance and most of them have been fully characterized the
function of many of them in the mechanism of stress tolerance is yet to be
investigated
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Ashraf M Breeding for salinity tolerance in plants Crit Rev Plant Sci 19941317 ndash42Ashraf M Akram NA Improving salinity tolerance of plants through conventional
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Ashraf M Foolad MR Roles of glycinebetaine and proline in improving plant abioticstress resistance Environ Exp Bot 200759206ndash16
Ashraf M Athar HR Harris PJC Kwon TR Some prospective strategies for improvingcrop salt tolerance Adv Agron 20089745-110
Asins MJ Present and future of quantitative trait locus analysis in plant breeding PlantBreed 2002121281ndash91
Babu RC Nguyen BD Chamarerk V Shanmugasundaram P Chezhian P Jeyaprakash P et alGeneticanalysis of droughtresistance in riceby molecular markers association betweensecondary traits and 1047297eld performance Crop Sci 2003431457ndash69
Babu RC Zhang J Blum A Ho THD Wu R Nguyen HT HVA1 a LEA gene from barleyconfers dehydration tolerance in transgenic rice( Oryzasativa L)via cell membraneprotection Plant Sci 2004166855ndash62
Badawi GH Kawano N Yamauchi Y Over-expression of ascorbate peroxidase in
tobacco chloroplasts enhances the tolerance to salt stress and water de1047297cit PhysiolPlant 2004121231ndash8Badu-Apraku B Yallou CG Registration of striga-resistant and drought tolerant tropical
early maize populations TZE-W Pop DT STR C4 and TZE-Y Pop DT STR C4 J PlantRegistr 20093(1)86ndash90
Badu-Apraku B Menkir A Kling JG Fakorede MAB Registration of 16 striga resistantearly maturing tropical maize inbred lines Crop Sci 2006461410ndash1
Baenziger PS Beecher B Graybosch RA Ibrahim AMH Baltensperger DD Nelson LA et alRegistration of lsquoNEO1643rsquo wheat J Plant Registr 20082(1)36ndash42
Baumlnziger M Setimela PS Hodson D Vivek B Breeding for improved drought tolerance inmaize adapted to southern Africa Proceedings of the 4th International Crop ScienceCongress Brisbane Australia Published on CDROM 2004 26 Sep ndash 1 Oct
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traits in the Mediterranean environment identi1047297ed in recombinant inbred lines of the cross lsquoArtarsquo H spontaneum 41-1 Theor Appl Genet 20031071215ndash25
Bergman JW Riveland NR Flynn CR Carlson GR Wichman DM Registration of lsquoMorlinrsquo
saf 1047298ower Crop Sci 2001411640Bernier J Kumar A Venuprasad R Spaner D Atlin G A large-effect QTL for
grain yield under reproductive-stage drought stress in upland rice Crop Sci200747507ndash18
BernierJ Kumar A SerrajR SpanerD Atlin G Review breeding uplandrice fordroughtresistance J Sci Food Agric 200888927ndash39
Bernier J Serraj R Kumar A Venuprasad R Impa S Gowdaa RPV et al The large-effectdrought-resistance QTL qtl121 increases water uptake in upland rice Field CropsRes 200911039ndash46
Bhatnagar-Mathur P ReddyDS Lavanya M Yamaguchi-Shinozaki K Sharma KK Stress-inducible expression of Arabidopsis thaliana DREB1A in transgenic peanut ( Arachishypogaea L) increases transpiration ef 1047297ciency under water-limiting conditionsPlant Cell Rep 2007262071ndash82
Bhatnagar-Mathur P Devi MJ Vadez V Sharma KK Differential antioxidative responses intransgenic peanut bear no relationship to their superior transpiration ef 1047297ciency underdrought stress J Plant Physiol 2009166(11)1207ndash17 doi101016jjplph200901001
Bidinger FR Serraj R Rizvi SMH Howarth C Yadav RS Hash CT Field evaluation of droughttolerance QTL effects on phenotype and adaptation in pearl millet [ Pennisetum glaucum(L) R Br] topcross hybrids Field Crops Res 200594(1)14ndash32
Bidinger FR Nepolean T Hash CT Yadav RS Howarth CJ Identi1047297cation of QTLs for grainyield of pearl millet (Pennisetum glaucum (L) R Br) in environments with variablemoisture during grain 1047297lling Crop Sci 200747969ndash80
Bowers JE Abbey C Anderson S Chang C Draye X A high-density geneticrecombination map of sequence-tagged sites for Sorghum as a framework forcomparative structural and evolutionary genomics of tropical grains and grassesGenetics 2003165367ndash86
Brick MA Ogg JB Singh SP Schwartz HF Johnson JJ Pastor-Corrales MA Registration of drought-tolerant rust-resistant high-yielding pinto bean germplasm lineCO46348 J Plant Registr 20082(2)120ndash4
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Busk PK Pages M Regulation of abscisic acid induced transcription Plant Mol Biol199837425ndash35
Carena MJ Wanner DW Development of genetically broad-based inbred lines of maizefor early-maturing (70-80RM) hybrids J Plant Registr 20093107ndash11
Cash SDBrucknerPL Wichman DMKephart KD Berg JEBoynerR et alRegistration of Willow Creek forage wheat J Plant Registr 20093(2)185ndash90
Cattivelli L Rizza F Badeck FW Mazzucotelli E Francia AMEM Mare AT et al Droughttolerance improvement in crop plants an integrated view from breeding to genomics
Field Crops Res 20081051-14Chen M Wang QY Cheng XG Xu ZS Li LC Ye XG et al GmDREB2 a soybean DRE-
binding transcription factor conferred drought and high-salt tolerance intransgenic plants Biochem Biophys Res Commun 2007353299ndash305
Cheng Z Targolli J Huang X Wu R Wheat LEA genes PMA80 and PMA1959 enhancedehydration tolerance of transgenic rice (Oryza sativa L) Mol Breed 20021071ndash82
ChoEK Hong ChB Over-expression of tobacco NtHSP70-1 contributes to drought-stresstolerance in plants Plant Cell Reports 200625349ndash58
Close TJ Dehydrins a commonality in the response of plants to dehydration and lowtemperature Physiol Plant 1997100291ndash6
Collins NC Tardieu F Tuberosa R Quantitative trait loci and crop performance underabiotic stress where do we stand Plant Physiol 2008147469ndash86
Concept Note Combining breeding and biotechnology to develop water ef 1047297cient maizefor Africa (WEMA) Afr Agric Technol Foundation
Courtois BShen L Petalcorin WCarandang S MauleonR LiZ Locating QTLs controllingconstitutive root traits in the rice population IAC 165-Co39 Euphytica 2003134335ndash45
Da-hong L Hui L Yang YL Ping-ping Z Jian-sheng L Down-regulated expression of RACK1geneby RNAinterference enhancesdroughttolerancein rice RiceSci 200916(1)14ndash20
Dalal M Tayal D Chinnusamy V Bansala KC Abiotic stress and ABA-inducible group 4 LEAfrom Brassicanapus plays a key rolein saltand droughttolerance J Biotechnol 2009139137ndash45
Ding Z Li S An X Liu X Qin H Wang D Transgenic expression of MYB15 confers enhancedsensitivity to abscisic acid and improved drought tolerance in Arabidopsis thaliana
J Genet Genomics 20093617ndash29Dubouzet JG Sakuma Y Ito Y Kasuga M Dubouzet EG Miura S et al OsDREB genes in
rice Oryza sativa L encode transcription activators that function in drought high-salt- and cold-responsive gene expression Plant J 200333751ndash63
El-Far IA Allan AY Responses of some wheat cultivars to sowing methods and droughtat different stages of growth Assuit J Agric Sci 199526(1)267ndash77
Eltayeb AE Kawano N Badawi GH Kaminaka H Sanekata T Shibahara T et alOverexpression of monodehydroascorbate reductase in transgenic tobacco confersenhanced tolerance to ozonesalt and polyethyleneglycol stresses Planta 2007225(5)1255ndash64
Falconer DS Introduction to quantitative genetics London New York Longman 1989Feng-ling FU Zhi-Lei F Shi-bing G Shu-feng Z Wan-chen L Evaluation and quantitative
inheritanceof several drought-relativetraits in maize Agric SciChina 20087(3)280ndash90Finkelstein R Gampala S Rock C Abscisic acid signaling in seeds and seedlings Plant
Cell 20021415ndash45Flowers TJ Improving crop salt tolerance J Exp Bot 200455307ndash19Fujita Y Fujita M Satoh R Maruyama K Parvez MM Seki M et al AREB1 is a transcription
Giraudat J Parcy F Bertauche N Gosti F Leung J Morris PC et al Current advances inabscisic acid action and signalling Plant Mol Biol 1994261557ndash77
Gorbalenya AE Koonin EV Helicases amino acid sequence comparisons and structurendashfunction relationships Curr Opin Struct Biol 19933419ndash29
Gosal SS Wani SH Kang MS Biotechnology and drought tolerance J Crop Improvement20092319ndash54
Gubis J Vaňkovaacute R Červenaacute V Draguacuteňovaacute M Hudcovicovaacute M Lichtnerovaacute H et alTransformedtobacco plants with increased toleranceto droughtSouthAfr J Bot200773505ndash11
GuoO Zhang J GaoQ Xing SLi F Wang WDrought tolerancethrough over-expressionof mono ubiquitin in transgenic tobacco J Plant Physiol 20081651745 ndash55
Haley SD Johnson JJ Peairs FB Quick JS Stromberger JA Clayshulte SR et al Registration of lsquoRipperrsquo wheat J Plant Registr 200711ndash6
Han SE Park SR Kwon HB Yi BY Lee GB Byun MO Genetic engineering of drought-resistant tobacco plants by introducingthe trehalose phosphorylase (TP) gene fromPleurotus sajor-caju Plant Cell Tissue Organ Cult 200582151ndash8
Harris K Klein R Mullet J Sorghum stay-green QTL individually reduces post-1047298oweringdrought-induced leaf senescence J Exp Bot 200758327ndash38
Hong BS Zong-Suo L Ming-An S LEA proteins in higher plants structure functiongene expression and regulation Colloids Surf B Biointerf 200545131ndash5
Howarth CJ Yadav RS Successful marker assisted selection for drought tolerance anddisease resistance in pearl milletIGER Innovations 2002
Humphreys MO Humphreys MW Breeding for stress resistance general principles InAshraf M Harris PJC editors Abiotic stresses plant resistance through breedingand molecular approaches 2005 p 19ndash46
Ingram J Bartels D The molecular basis of dehydration tolerance in plants Ann RevPlant Physiol Plant Mol Biol 199647377ndash403
Jaglo-Ottosen KR Gilmour SJ Zarka DG Schabenberger O Thomashow MF ArabidopsisCBF1 overexpression induces COR genes and enhances freezing tolerance Science1998280104ndash6
Jang IC Oh SJ Seo JS Choi WB Song SI Kim CH et al Expression of a bifunctional fusion of the Escherichia coli genes for trehalose-6-phosphate synthase and trehalose-6-phosphate phosphatase in transgenic rice plants increases trehalose accumulationandabiotic stresstolerancewithoutstuntinggrowth Plant Physiol2003131516ndash24
Jeanneau M Gerentes D Foueillassar X Zivy M Vidal J Toppan A et al Improvement of drought tolerance in maize towards the functional validation of the Zm-Asr1 gene andincrease of water use ef 1047297ciency by over-expressing C4-PEPC Biochimie 2002841127ndash35
Jenson KB Palazzo AJ Waldron BL Bushman BS Registration of lsquoFirstStrikersquo Slenderwheatgrass J Plant Registr 2007124ndash5
Joung-youn K Hyung-in C Min-young I Soo-young K Arabidopsis basic leucine zipperproteins that mediate stress-responsive abscisic acid signaling Plant Cell 20024343ndash57
Juskiw PE Helm JH Oro M Nyachiro JM Salmon DF Registration of lsquoBentleyrsquo barley
J Plant Registr 20093(2)119ndash23Kalamaki MS Alexandrou D Lazari D Merkouropoulos G Fotopoulos V Pateraki I et al
Over-expression of a tomato N-acetyl-L -glutamate synthase gene (SlNAGS1) in Arabidopsis thaliana results in high ornithine levels and increased tolerance in saltand drought stresses J Exp Bot 200960(6)1859ndash71
Kamoshita A Babu CR Boopathi NM Fukai S Phenotypic and genotypic analysis of drought-resistance traits for development of rice cultivars adapted to rainfedenvironments Field Crops Res 2008109(103)1-23 doi101016jfcr200806010
KarakasB Ozias-AkinsP Stushnoff C SuefferheldM Rieger M Salinityand drought toleranceof mannitol-accumulating transgenic tobacco Plant Cell Environ 199720609ndash16
Karim S Aronsson H Ericson H Pirhonen M Leyman B Welin B et al Improved droughttolerance without undesired side effects in transgenic plants producing trehalose PlantMol Biol 200764371ndash86
Kasuga M Liu Q Miura S Yamaguchi-Shinozaki K Shinozaki K Improving plant droughtsaltand freezing tolerance by gene transfer of a single stress-inducible transcription factorNat Biotechnol 199917287ndash91
Kindiger M Gaub H Hasegawac M Katsurab Y Ueyamad K Gotob S et al Registrationof lsquoNanryorsquo tall fescue Crop Sci 2006461815ndash6
Kong I Dong J HartGE Characteristics linkage mappositionsand allelic differentiationof Sorghum bicolour (L) Moench by DNA simple-sequence repeats (SSRs) TheorAppl Genet 2000101438ndash48
Kumar R Venuprasad R Atlin GN Genetic analysis of rainfed lowland rice droughttolerance under naturally-occurring stress in eastern India heritability and QTL effects Field Crops Res 200710342ndash52
La1047297tte HR Price AH Courtois B Yield response to water de1047297cit in an upland ricemapping population associations among traits and genetic markers Theor ApplGenet 20041091237ndash46
Lal S Gulyani V Khurana P Overexpressionof HVA1 gene frombarley generates tolerance tosalinity and water stress in transgenic mulberry (Morus indica) Transgenic Res 200817
(4)651ndash
63Lanceras J Pantuwan G Jongdee B Toojinda T Quantitative trait loci associated withdrought tolerance at reproductive stage in rice Plant Physiol 2004135384ndash99
Levi A Ovnat L Paterson AH Saranga Y Photosynthesis of cotton near-isogenic linesintrogressed with QTLs for productivity and drought related traits Plant Sci 2009a17788ndash96
Levi A Paterson AH Barak V Yakir D Wang B Chee PW et al Field evaluation of cottonnear-isogenic lines introgressed with QTLs for productivity and drought relatedtraits Mol Breed 2009b23179ndash95
Liu Q Kasuga M Sakuma Y Abe H Miura S Yamaguchi- Shinozaki K et al Twotranscription factors DREB1 and DREB2 with an EREBPAP2 DNA binding domainseparate two cellular signal transduction pathways in drought-and low-temper-ature-responsive gene expression respectively in Arabidopsis Plant Cell 1998101391ndash406
Liu X Hua X Guo J Qi D Wang L Liu Z et al Enhanced tolerance to drought stress intransgenic tobacco plants overexpressing Biotechnol Lett 2008301275ndash80
Liu X Wanga Z Wanga L Wua R Phillips J Deng X LEA 4 group genes from theresurrection plant Boea hygrometrica confer dehydration tolerance in transgenictobacco Plant Sci 200917690ndash8
Luchi S Kobayashi M Taji T Naramoto M Seki M Kato T et al Regulation of droughttolerance by gene manipulation of 9-cis-epoxycarotenoid dioxygenase a keyenzyme in abscisic acid biosynthesis in Arabidopsis Plant J 200127325ndash33
Luo Y Liu YB Dong YX Gao XQ Zhang YS Expression of a putative alfalfa helicase increasestolerance to abioticstress in Arabidopsis by enhancing the capacities for ROS scavengingand osmotic adjustment J Plant Physiol 2009166385ndash94
MacLean JLDaweDC Hardy B Hettel GPRiceAlmanac sourcebook for themostimportanteconomic activity on Earth3rd ed Wallingford England CABI Publishing 2002
Mansour MMF Nitrogen containing compounds and adaptation of plants to salinitystress Biol Plant 200043491ndash500
McKersie BD Bowley SR Harjanto E Leprince O Water-de1047297cit tolerance and 1047297eldperformance of transgenic alfalfa overexpressing superoxide dismutase Plant Physiol19961111177ndash81
McKersie BD Murnaghan J Bowley SR Manipulating freezing tolerance in transgenicplants Acta Physiol Plant 199719485ndash95
Miranda JA Avonce N Suaacuterez R Thevelein JM Dijck PV Iturriaga G A bifunctional TPS-TPP enzyme from yeast confers tolerance to multiple and extreme abiotic-stressconditions in transgenic Arabidopsis Planta 2007226(6)1411ndash21
Mohammadi M Taleei A Zeinali H Naghavi MR Ceccarelli S Grando Baum M QTL analysis for phenologic traits in doubled haploid population of barley Int J AgricBiol 20057(5)820ndash3
Mohanty A Kathuria H Ferjani A Sakamoto A Mohanty P Murata N et al Transgenicsof an elite indica rice variety Pusa Basmati 1 harbouring the codA gene are highlytolerant to salt stress Theor Appl Genet 200210651ndash7
Noaman MM El Sayad AA Asaad FA El Sherbini AM El Bawab AO El Moselhi MA et alRegistration of lsquoGiza 126rsquo barley Crop Sci 199535(6)1710
Noaman MM Ahmed IA El-Sayed AA Abo-El-Enin RA El-Gamal AS El-Sherbiny AM et alRegistration of lsquoGiza 2000rsquo drought-tolerant six-rowed barley for rainfed and newreclaimed areas in Egypt Crop Sci 200747440
Obert DE Evans CP Wesenberg DM Windes JM Erickson CA Jackson EW et alRegistration of lsquoLenetahrsquo spring barley J Plant Registr 20082(2)85ndash7
Oh SJ Jeong JS Kim EH Yi NR Yi SI Jang IC et al Matrix attachment region fromthe chicken lysozyme locus reduces variability in transgene expression andconfers copy number-dependence in transgenic rice plants Plant Cell Rep20054145ndash54
Oh SJ Kim YS Kwon C Park HK Jeong JS Kim JK Overexpression of the transcriptionfactor AP37 in rice improves grain yield under drought conditions Plant Physiol
20092191ndash
200 doi101104pp109137554Owttrim GW RNA helicases and abiotic stress Nucleic AcidsRes 200634(11)3220ndash30Park BJ Liu Z Kanno A Kameya T Increased tolerance to salt and water de1047297cit stress in
transgenic lettuce (Lactuca sativa L) by constitutive expression of LEA PlantGrowth Regul 2005a45165ndash71
Park BJ Liu Z Kanno A Kameya T Genetic improvement of Chinese cabbage for salt anddroughttolerance by constitutiveexpression of a B napus LEAgenePlantSci 2005b169553ndash8
Pastori GM Foyer CH Common componentsnetworks and pathways of crosstoleranceto stress The central role of ldquoredoxrdquo and abscisic acid-mediated controls PlantPhysiol 2002129460ndash8
PellegrineschiA ReynoldsM PachecoM Brito RMAlmeraya R Yamaguchi-ShinozakiK et alStress-induced expression in wheat of the Arabidopsis thaliana DREB1Agenedelayswaterstress symptoms under greenhouse conditions Genome 200447493ndash500
Perl A Perl-Treves R Galili S Aviv D Shalgi E Malkin S et al Enhanced oxidative-stressdefense in transgenic potato overexpressing tomato Cu Zn superoxide dismutaseTheor Appl Genet 199385568ndash76
Podlich DW Winkler CR Cooper M Mapping as you go an effective approach formarker-assisted selection of complex traits Crop Sci 2004441560ndash71
182 M Ashraf Biotechnology Advances 28 (2010) 169ndash183
Pratt RC Casey MA Registration of maize germplasm line Oh605 Crop Sci 2006461004ndash5Qin F Kakimoto M Sakuma Y Maruyama K Osakabe Y Tran LSP et al Regulation and
functional analysis of ZmDREB2A in response to drought and heat stress in Zea mays LPlant J 20075054ndash69
Quan R Shang M Zhang H Zhao Y Zhang J Engineering of enhanced glycinebetainesynthesis improves drought tolerance in maize Plant Biotechnol J 20042477ndash86
Quarrie SA Gulli M Calestani C Steed A Marmiroli N Location of a gene regulatingdrought-induced abscisic acid production on the long arm of chromosome 5A of wheat Theor Appl Genet 199489794ndash800
QuickJS Stromberger JA Clayshulte S Clifford B Johnson JJ Peairs FB et al Registrationof lsquoPrairie Redrsquo wheat Crop Sci 2001411362ndash3
RajaramS Prospects andpromise ofwheatbreedingin the21stcentury Euphytica 20011193-15Rajaram S Role of conventional plant breeding and biotechnology in future wheat
production Turk J Agric Forest 200529105ndash11Rami JF Dufour P Trouche G Fliedel G Mestres C Davrieux F et al Quantitative trait
loci for grain quality productivity morphological and agronomical traits insorghum (Sorghum bicolor L Moench) Theor Appl Genet 199897605ndash16
Rassaa N Salahb HBH Latiri K Thermal responses of durum wheat Triticum durum toearly water stress consequence on leaf and 1047298ower development Plant Biol Pathol2008331(5)363ndash71
Reddy LJ Nigam SN Rao RCN Reddy NS Registration of ICGV 87354 peanut germplasmwith drought tolerance and rust resistance Crop Sci 200141274ndash5
Ribaut JM Ragot M Marker-assisted selection to improve drought adaptation in maize thebackcross approach perspectives limitations and alternatives J Exp Bot 200658351ndash60
Robin S Pathan MS Courtois B La1047297tte R Carandang S Lanceras S et al Mappingosmotic adjustment in an advanced back-cross inbred population of rice Theor
Appl Genet 2003107(7)1288ndash96Romero C Belles JM Vaya JL Serrano R Culianez-Macia FA Expression of the yeast
trehalose-6-phosphate synthase gene in transgenic tobacco plants pleiotropicphenotypes include drought tolerance Planta 1997201293ndash7
Ronde JAD Cress WA Krugerd GHJ Strasserd RJ Van Staden J Photosynthetic responseof transgenic soybean plants containing an Arabidopsis P5CR gene during heat anddrought stress J Plant Physiol 20041611211ndash24
Salvi S Tuberosa R To clone or not to clone plant QTLs present and future challengesTrends Plant Sci 200510297ndash304
Sanchez AC Subudhi PK Rosenow DT Nguyen HT Mapping QTLs associated with droughtresistance in sorghum (Sorghum bicolor L Moench) Plant Mol Biol 200248713ndash26
Saranga Y Menz M Jiang CX Wright RJ Yakir D Paterson AH Genomic dissection of genotypetimesenvironment interactions conferring adaptation of cotton to aridconditions Genome Res 2001111988ndash95
Sari-Gorla M Krajewski P Di Fonzo N Villa M Frova C Genetic analysis of droughttolerance in maize by molecular markers II Plant height and 1047298owering Theor ApplGenet 199999289ndash95
Seki M Narusaka M Ishida J Nanjo T Fujita M Oono Y et al Monitoring the expressionpro1047297les of 7000 Arabidopsis genes under drought cold and high-salinity stressesusing a full-length cDNA microarray Plant J 200231279ndash92
Serraj R Sinclair TR Osmolyte accumulation can it really increase crop yield underdrought conditions Plant Cell Environ 200225333ndash41
Serraj R Krishnamurthy L Kashiwagi J Kumar J Chandra S Crouch JH Variation in roottraits of chickpea (Cicer arietinum L) grown under terminal drought Field CropsRes 200488115ndash27
Serraj R Hash CT Rizvi MHS Sharma A Yadav RS Bidinger FR Recent advances in marker-assisted selection for drought tolerance in pearl millet Plant Prod Sci 20058(3)334ndash7
Shen YG Du BX Zhang WK Zhang JS Chen SY AhCMO regulated by stresses in Atriplexhortensis can improve drought tolerance in transgenic tobacco Theor Appl Genet2002105815ndash21
Shinozaki K Yamaguchi-Shinozaki K Molecular responses to dehydration and lowtemperature differences and cross-talk between two stress signaling pathwaysCurr Opin Plant Biol 20003217ndash23
Singh KB Omar M Saxena MC Johansen C Registration of FLIP 87-59C a drought-tolerant chickpea germplasm line Crop Sci 199636(2)1ndash2
Singh SP Teran H Gutierrez JA Registration of SEA 5 and SEA 13 drought tolerant drybean germplasm Crop Sci 200141276ndash7
Singh SP Teran H Lema M Schwartz HF Miklas PN Registration of white moldresistant dry bean germplasm line A 195 J Plant Registr 2007162ndash3
Sivamani E Bahieldin A Wraith JM Improved biomass productivity and water useef 1047297ciency under water de1047297cit conditions in transgenic wheat constitutivelyexpressing the barley HVA1 gene Plant Sci 20001551ndash9
Skovmand B Reynolds MP DeLacy IH Searching genetic resources for physiologicaltraits with potential for increasing yield In Reynolds MP Ortiz-Monasterio IMcNab A editors Application of Physiology in Wheat Breeding 2001 p 17ndash28
Steele K Novel upland rice variety bred using marker-assisted selection and client-oriented breeding released in Jharkhand India Bangor University 2009
Steele KA Price AH Shashidar HE Witcombe JR Marker-assistedselection to introgressrice QTLs controlling root traitsinto an Indianupland rice variety Theor Appl Genet2006112208ndash21
Steele KA Virk DS Kumar R Prasad SC Witcombe JR Field evaluation of upland ricelines selected for QTLs controlling root traits Field Crops Res 2007101180 ndash6
Sunkar R Kapoor A Zhu JK Post transcriptional induction of two CuZn superoxidedismutase genes in Arabidopsis is mediated by down regulation of miR398 andimportant for oxidative stress tolerance Plant Cell 2006182051ndash65
Talame V Sanguineti MC Chiapparino E Bahri H Ben Salem M Forster BP et alIdenti1047297cation of Hordeum spontaneum QTL alleles improving 1047297eld performance of barley grown under rainfed conditions Ann Appl Bot 2004144309ndash20
Tanner NKCordinO BanroquesJ DoereM Linder PThe Q Motif a newlyidenti1047297ed motif in DEAD box helicases may regulate ATP binding and hydrolysis Mol Cell 200311127ndash38
Taramino G Tarchini R Ferrario S Lee M Pe ME Characterization and mapping of simplesequence repeats (SSRs) in Sorghum bicolor Theor Appl Genet 19979566ndash72
Teulat B Monneveux P Wery J Borriegraves C Souyris I Charrier A et al Relationshipsbetween relative water content andgrowth parameters in barleya QTLstudy NewPhytol 199713799-107
Thi Lang N Chi Buu B Fine mapping for drought tolerance in rice ( Oryza sativa L)Omonrice 2008169-15Till BJ Comai L Henikoff S Tillering and ecotillering for crop improvement In
Varshney RK Tuberosa R editors Genomics-assisted crop improvement genomicsapproaches and platforms vol 1 2007 p 333ndash50
Tondelli A Francia E Barabaschi D Aprile A Skinner JS Stockinger EJ et al Mappingregulatory genes as candidates for cold and drought stress tolerance in barleyTheor Appl Genet 2006112445ndash54
Tuberosa R Salvi S Genomics approaches to improve drought tolerance in cropsTrends Plant Sci 200611405ndash12
Valkoun JJ Wheat pre-breeding using wild progenitors Euphytica 200111917ndash23Vashisht AA Tuteja N Stress responsive DEAD-box helicases a new pathway to
engineer plant stress tolerance J Photochem Photobiol B Biol 200684150 ndash60Vendruscolo ECG Schuster I Pileggi M Scapim CA Molinari HBC Marur CJ et al Stress-
induced synthesis of proline confers tolerance to water de1047297cit in transgenic wheat J Plant Physiol 20071641367ndash76
Vienne D Leonardi A Damerval C Zivy M Genetics of proteome variation for QTL characterization application to drought stress responses in maize J Exp Bot 199950303ndash9
Villareal RL Mujeeb-Kazi A Rajaram S Toro ED Morphological variability in somesynthetic hexaploid wheats derived from Triticum turgidum times T tauschii J GenetBreed 1994487-16
Vinh NT Paterson AH Genome mapping and its implication for stress resistance inplants In Ashraf M Harris PJC editors Abiotic stresses plant resistance throughbreeding and molecular approaches 2005
Vinocur B Altman A Recent advances in engineering plant tolerance to abiotic stressachievements and limitations Curr Opin Biotechnol 200516123ndash32
Wang YJ Hao YJ Zhang ZG Chen T Zhang JS Chen SY Isolation of trehalose-6-phosphate phosphatase gene from tobacco and its functional analysis in yeast cells
J Plant Physiol 2005162215ndash23Wu R Garg A Engineering rice plants with trehalose-producing genes improves
tolerance to drought salt and low temperature ISB News Report 2003Xiao B Huang Y Tang N Xiong L Overexpression of LEA gene in rice improves drought
resistance under 1047297eld conditions Theor Appl Genet 200711535ndash46Xinglai P Sangang X Qiannying P Yinhong S Registration of lsquo Jinmai 50rsquo wheat Crop Sci
200646983ndash5Xiong L Lee H Ishitani M Zhu JK Regulation of osmotic stress-responsive gene
expression by the LOS6ABA1 locus in Arabidopsis J Biol Chem 20022778588ndash96Xu GW Magill CW Shertz KF Hart GE A RFLP linkage map of Sorghum bicolor (L)
Moench Theor Appl Genet 199489139ndash45Xu D Duan X Wang B Hong BHo THD Wu RExpression of a late embryogenesis abundant
protein gene HVA1 from barley confers tolerance to water de1047297cit and salt stress intransgenic rice Plant Physiol 1996110249ndash57
Yadav RS Hash CT Bidinger FR Devos KM Howarth CJ Genomic regions associatedwith grain yield and aspects of post-1047298owering drought tolerance in pearl milletacross stress environments and testers background Euphytica 2004136265ndash77
YamadaM MorishitaH Urano K Shiozaki N Yamaguchi-Shinozaki K ShinozakiK et alEffects of free proline accumulation in petunias under drought stress J Exp Bot2005561975ndash81
Yang WJ Rich PJ Axtell JD Wood KV Bonham CC Ejeta G et al Genotypic variation forglycinebetaine in sorghum Crop Sci 200343162ndash9
Yan-Ying QU Ping MU Xue-Qin L Yu-Xiu T Feng W Hong-Liang Z et al QTL mappingand correlations between leaf water potential and drought resistance in rice underupland and lowland environments Acta Agron Sin 200834(2)198ndash206
Ye XDWu XLZhaoH Frehner MNoumlsberger J Potrykus Iet al Altered fructan accumulationin transgenic Lolium multi 1047298orum plants expressing a Bacillus subtilis sacB gene Plant Cell
Rep 200120205ndash
12Zaharieva M Gaulin E Havaux M Acevedo E Monneveux P Drought and heat responses in
the wild wheat relative Aegilops geniculata Roth potential interest for wheatimprovement Crop Sci 2001411321ndash9
Zhang J Zheng HGAartiA Pantuwan G NguyenTT Tripathy JNet alLocatinggenomicregions associated with components of drought resistance in rice comparativemapping within and across species Theor Appl Genet 200110319ndash29
Zhang GH Su Q An LJ Wu S Characterization and expression of a vacuolar Na+H+
antiporter gene from the monocot halophyte Aeluropus littoralis Plant PhysiolBiochem 200846117ndash26
Zhao J QTLs for oil content and their relationships to other agronomic traits in anEuropeantimesChinese oilseed rape population Germany Diss Grorg-Agust Univer-sity of Goettingen 2002
Zhao J Ren W Zhi D Wang L Xia G Arabidopsis DREB1ACBF3 bestowed transgenic tallfescue increased tolerance to drought stress Plant Cell Rep 2007261521ndash8
Zhou W Li Y Zhao BC Ge RC Shen YZ Wang G Huang ZJ Over-expression of TaSTRG gene improves salt and drought tolerance in rice J Plant Physiol 200915(166)1660ndash70
Dalal M Tayal D Chinnusamy V Bansala KC Abiotic stress and ABA-inducible group 4 LEAfrom Brassicanapus plays a key rolein saltand droughttolerance J Biotechnol 2009139137ndash45
Ding Z Li S An X Liu X Qin H Wang D Transgenic expression of MYB15 confers enhancedsensitivity to abscisic acid and improved drought tolerance in Arabidopsis thaliana
J Genet Genomics 20093617ndash29Dubouzet JG Sakuma Y Ito Y Kasuga M Dubouzet EG Miura S et al OsDREB genes in
rice Oryza sativa L encode transcription activators that function in drought high-salt- and cold-responsive gene expression Plant J 200333751ndash63
El-Far IA Allan AY Responses of some wheat cultivars to sowing methods and droughtat different stages of growth Assuit J Agric Sci 199526(1)267ndash77
Eltayeb AE Kawano N Badawi GH Kaminaka H Sanekata T Shibahara T et alOverexpression of monodehydroascorbate reductase in transgenic tobacco confersenhanced tolerance to ozonesalt and polyethyleneglycol stresses Planta 2007225(5)1255ndash64
Falconer DS Introduction to quantitative genetics London New York Longman 1989Feng-ling FU Zhi-Lei F Shi-bing G Shu-feng Z Wan-chen L Evaluation and quantitative
inheritanceof several drought-relativetraits in maize Agric SciChina 20087(3)280ndash90Finkelstein R Gampala S Rock C Abscisic acid signaling in seeds and seedlings Plant
Cell 20021415ndash45Flowers TJ Improving crop salt tolerance J Exp Bot 200455307ndash19Fujita Y Fujita M Satoh R Maruyama K Parvez MM Seki M et al AREB1 is a transcription
Giraudat J Parcy F Bertauche N Gosti F Leung J Morris PC et al Current advances inabscisic acid action and signalling Plant Mol Biol 1994261557ndash77
Gorbalenya AE Koonin EV Helicases amino acid sequence comparisons and structurendashfunction relationships Curr Opin Struct Biol 19933419ndash29
Gosal SS Wani SH Kang MS Biotechnology and drought tolerance J Crop Improvement20092319ndash54
Gubis J Vaňkovaacute R Červenaacute V Draguacuteňovaacute M Hudcovicovaacute M Lichtnerovaacute H et alTransformedtobacco plants with increased toleranceto droughtSouthAfr J Bot200773505ndash11
GuoO Zhang J GaoQ Xing SLi F Wang WDrought tolerancethrough over-expressionof mono ubiquitin in transgenic tobacco J Plant Physiol 20081651745 ndash55
Haley SD Johnson JJ Peairs FB Quick JS Stromberger JA Clayshulte SR et al Registration of lsquoRipperrsquo wheat J Plant Registr 200711ndash6
Han SE Park SR Kwon HB Yi BY Lee GB Byun MO Genetic engineering of drought-resistant tobacco plants by introducingthe trehalose phosphorylase (TP) gene fromPleurotus sajor-caju Plant Cell Tissue Organ Cult 200582151ndash8
Harris K Klein R Mullet J Sorghum stay-green QTL individually reduces post-1047298oweringdrought-induced leaf senescence J Exp Bot 200758327ndash38
Hong BS Zong-Suo L Ming-An S LEA proteins in higher plants structure functiongene expression and regulation Colloids Surf B Biointerf 200545131ndash5
Howarth CJ Yadav RS Successful marker assisted selection for drought tolerance anddisease resistance in pearl milletIGER Innovations 2002
Humphreys MO Humphreys MW Breeding for stress resistance general principles InAshraf M Harris PJC editors Abiotic stresses plant resistance through breedingand molecular approaches 2005 p 19ndash46
Ingram J Bartels D The molecular basis of dehydration tolerance in plants Ann RevPlant Physiol Plant Mol Biol 199647377ndash403
Jaglo-Ottosen KR Gilmour SJ Zarka DG Schabenberger O Thomashow MF ArabidopsisCBF1 overexpression induces COR genes and enhances freezing tolerance Science1998280104ndash6
Jang IC Oh SJ Seo JS Choi WB Song SI Kim CH et al Expression of a bifunctional fusion of the Escherichia coli genes for trehalose-6-phosphate synthase and trehalose-6-phosphate phosphatase in transgenic rice plants increases trehalose accumulationandabiotic stresstolerancewithoutstuntinggrowth Plant Physiol2003131516ndash24
Jeanneau M Gerentes D Foueillassar X Zivy M Vidal J Toppan A et al Improvement of drought tolerance in maize towards the functional validation of the Zm-Asr1 gene andincrease of water use ef 1047297ciency by over-expressing C4-PEPC Biochimie 2002841127ndash35
Jenson KB Palazzo AJ Waldron BL Bushman BS Registration of lsquoFirstStrikersquo Slenderwheatgrass J Plant Registr 2007124ndash5
Joung-youn K Hyung-in C Min-young I Soo-young K Arabidopsis basic leucine zipperproteins that mediate stress-responsive abscisic acid signaling Plant Cell 20024343ndash57
Juskiw PE Helm JH Oro M Nyachiro JM Salmon DF Registration of lsquoBentleyrsquo barley
J Plant Registr 20093(2)119ndash23Kalamaki MS Alexandrou D Lazari D Merkouropoulos G Fotopoulos V Pateraki I et al
Over-expression of a tomato N-acetyl-L -glutamate synthase gene (SlNAGS1) in Arabidopsis thaliana results in high ornithine levels and increased tolerance in saltand drought stresses J Exp Bot 200960(6)1859ndash71
Kamoshita A Babu CR Boopathi NM Fukai S Phenotypic and genotypic analysis of drought-resistance traits for development of rice cultivars adapted to rainfedenvironments Field Crops Res 2008109(103)1-23 doi101016jfcr200806010
KarakasB Ozias-AkinsP Stushnoff C SuefferheldM Rieger M Salinityand drought toleranceof mannitol-accumulating transgenic tobacco Plant Cell Environ 199720609ndash16
Karim S Aronsson H Ericson H Pirhonen M Leyman B Welin B et al Improved droughttolerance without undesired side effects in transgenic plants producing trehalose PlantMol Biol 200764371ndash86
Kasuga M Liu Q Miura S Yamaguchi-Shinozaki K Shinozaki K Improving plant droughtsaltand freezing tolerance by gene transfer of a single stress-inducible transcription factorNat Biotechnol 199917287ndash91
Kindiger M Gaub H Hasegawac M Katsurab Y Ueyamad K Gotob S et al Registrationof lsquoNanryorsquo tall fescue Crop Sci 2006461815ndash6
Kong I Dong J HartGE Characteristics linkage mappositionsand allelic differentiationof Sorghum bicolour (L) Moench by DNA simple-sequence repeats (SSRs) TheorAppl Genet 2000101438ndash48
Kumar R Venuprasad R Atlin GN Genetic analysis of rainfed lowland rice droughttolerance under naturally-occurring stress in eastern India heritability and QTL effects Field Crops Res 200710342ndash52
La1047297tte HR Price AH Courtois B Yield response to water de1047297cit in an upland ricemapping population associations among traits and genetic markers Theor ApplGenet 20041091237ndash46
Lal S Gulyani V Khurana P Overexpressionof HVA1 gene frombarley generates tolerance tosalinity and water stress in transgenic mulberry (Morus indica) Transgenic Res 200817
(4)651ndash
63Lanceras J Pantuwan G Jongdee B Toojinda T Quantitative trait loci associated withdrought tolerance at reproductive stage in rice Plant Physiol 2004135384ndash99
Levi A Ovnat L Paterson AH Saranga Y Photosynthesis of cotton near-isogenic linesintrogressed with QTLs for productivity and drought related traits Plant Sci 2009a17788ndash96
Levi A Paterson AH Barak V Yakir D Wang B Chee PW et al Field evaluation of cottonnear-isogenic lines introgressed with QTLs for productivity and drought relatedtraits Mol Breed 2009b23179ndash95
Liu Q Kasuga M Sakuma Y Abe H Miura S Yamaguchi- Shinozaki K et al Twotranscription factors DREB1 and DREB2 with an EREBPAP2 DNA binding domainseparate two cellular signal transduction pathways in drought-and low-temper-ature-responsive gene expression respectively in Arabidopsis Plant Cell 1998101391ndash406
Liu X Hua X Guo J Qi D Wang L Liu Z et al Enhanced tolerance to drought stress intransgenic tobacco plants overexpressing Biotechnol Lett 2008301275ndash80
Liu X Wanga Z Wanga L Wua R Phillips J Deng X LEA 4 group genes from theresurrection plant Boea hygrometrica confer dehydration tolerance in transgenictobacco Plant Sci 200917690ndash8
Luchi S Kobayashi M Taji T Naramoto M Seki M Kato T et al Regulation of droughttolerance by gene manipulation of 9-cis-epoxycarotenoid dioxygenase a keyenzyme in abscisic acid biosynthesis in Arabidopsis Plant J 200127325ndash33
Luo Y Liu YB Dong YX Gao XQ Zhang YS Expression of a putative alfalfa helicase increasestolerance to abioticstress in Arabidopsis by enhancing the capacities for ROS scavengingand osmotic adjustment J Plant Physiol 2009166385ndash94
MacLean JLDaweDC Hardy B Hettel GPRiceAlmanac sourcebook for themostimportanteconomic activity on Earth3rd ed Wallingford England CABI Publishing 2002
Mansour MMF Nitrogen containing compounds and adaptation of plants to salinitystress Biol Plant 200043491ndash500
McKersie BD Bowley SR Harjanto E Leprince O Water-de1047297cit tolerance and 1047297eldperformance of transgenic alfalfa overexpressing superoxide dismutase Plant Physiol19961111177ndash81
McKersie BD Murnaghan J Bowley SR Manipulating freezing tolerance in transgenicplants Acta Physiol Plant 199719485ndash95
Miranda JA Avonce N Suaacuterez R Thevelein JM Dijck PV Iturriaga G A bifunctional TPS-TPP enzyme from yeast confers tolerance to multiple and extreme abiotic-stressconditions in transgenic Arabidopsis Planta 2007226(6)1411ndash21
Mohammadi M Taleei A Zeinali H Naghavi MR Ceccarelli S Grando Baum M QTL analysis for phenologic traits in doubled haploid population of barley Int J AgricBiol 20057(5)820ndash3
Mohanty A Kathuria H Ferjani A Sakamoto A Mohanty P Murata N et al Transgenicsof an elite indica rice variety Pusa Basmati 1 harbouring the codA gene are highlytolerant to salt stress Theor Appl Genet 200210651ndash7
Noaman MM El Sayad AA Asaad FA El Sherbini AM El Bawab AO El Moselhi MA et alRegistration of lsquoGiza 126rsquo barley Crop Sci 199535(6)1710
Noaman MM Ahmed IA El-Sayed AA Abo-El-Enin RA El-Gamal AS El-Sherbiny AM et alRegistration of lsquoGiza 2000rsquo drought-tolerant six-rowed barley for rainfed and newreclaimed areas in Egypt Crop Sci 200747440
Obert DE Evans CP Wesenberg DM Windes JM Erickson CA Jackson EW et alRegistration of lsquoLenetahrsquo spring barley J Plant Registr 20082(2)85ndash7
Oh SJ Jeong JS Kim EH Yi NR Yi SI Jang IC et al Matrix attachment region fromthe chicken lysozyme locus reduces variability in transgene expression andconfers copy number-dependence in transgenic rice plants Plant Cell Rep20054145ndash54
Oh SJ Kim YS Kwon C Park HK Jeong JS Kim JK Overexpression of the transcriptionfactor AP37 in rice improves grain yield under drought conditions Plant Physiol
20092191ndash
200 doi101104pp109137554Owttrim GW RNA helicases and abiotic stress Nucleic AcidsRes 200634(11)3220ndash30Park BJ Liu Z Kanno A Kameya T Increased tolerance to salt and water de1047297cit stress in
transgenic lettuce (Lactuca sativa L) by constitutive expression of LEA PlantGrowth Regul 2005a45165ndash71
Park BJ Liu Z Kanno A Kameya T Genetic improvement of Chinese cabbage for salt anddroughttolerance by constitutiveexpression of a B napus LEAgenePlantSci 2005b169553ndash8
Pastori GM Foyer CH Common componentsnetworks and pathways of crosstoleranceto stress The central role of ldquoredoxrdquo and abscisic acid-mediated controls PlantPhysiol 2002129460ndash8
PellegrineschiA ReynoldsM PachecoM Brito RMAlmeraya R Yamaguchi-ShinozakiK et alStress-induced expression in wheat of the Arabidopsis thaliana DREB1Agenedelayswaterstress symptoms under greenhouse conditions Genome 200447493ndash500
Perl A Perl-Treves R Galili S Aviv D Shalgi E Malkin S et al Enhanced oxidative-stressdefense in transgenic potato overexpressing tomato Cu Zn superoxide dismutaseTheor Appl Genet 199385568ndash76
Podlich DW Winkler CR Cooper M Mapping as you go an effective approach formarker-assisted selection of complex traits Crop Sci 2004441560ndash71
182 M Ashraf Biotechnology Advances 28 (2010) 169ndash183
Pratt RC Casey MA Registration of maize germplasm line Oh605 Crop Sci 2006461004ndash5Qin F Kakimoto M Sakuma Y Maruyama K Osakabe Y Tran LSP et al Regulation and
functional analysis of ZmDREB2A in response to drought and heat stress in Zea mays LPlant J 20075054ndash69
Quan R Shang M Zhang H Zhao Y Zhang J Engineering of enhanced glycinebetainesynthesis improves drought tolerance in maize Plant Biotechnol J 20042477ndash86
Quarrie SA Gulli M Calestani C Steed A Marmiroli N Location of a gene regulatingdrought-induced abscisic acid production on the long arm of chromosome 5A of wheat Theor Appl Genet 199489794ndash800
QuickJS Stromberger JA Clayshulte S Clifford B Johnson JJ Peairs FB et al Registrationof lsquoPrairie Redrsquo wheat Crop Sci 2001411362ndash3
RajaramS Prospects andpromise ofwheatbreedingin the21stcentury Euphytica 20011193-15Rajaram S Role of conventional plant breeding and biotechnology in future wheat
production Turk J Agric Forest 200529105ndash11Rami JF Dufour P Trouche G Fliedel G Mestres C Davrieux F et al Quantitative trait
loci for grain quality productivity morphological and agronomical traits insorghum (Sorghum bicolor L Moench) Theor Appl Genet 199897605ndash16
Rassaa N Salahb HBH Latiri K Thermal responses of durum wheat Triticum durum toearly water stress consequence on leaf and 1047298ower development Plant Biol Pathol2008331(5)363ndash71
Reddy LJ Nigam SN Rao RCN Reddy NS Registration of ICGV 87354 peanut germplasmwith drought tolerance and rust resistance Crop Sci 200141274ndash5
Ribaut JM Ragot M Marker-assisted selection to improve drought adaptation in maize thebackcross approach perspectives limitations and alternatives J Exp Bot 200658351ndash60
Robin S Pathan MS Courtois B La1047297tte R Carandang S Lanceras S et al Mappingosmotic adjustment in an advanced back-cross inbred population of rice Theor
Appl Genet 2003107(7)1288ndash96Romero C Belles JM Vaya JL Serrano R Culianez-Macia FA Expression of the yeast
trehalose-6-phosphate synthase gene in transgenic tobacco plants pleiotropicphenotypes include drought tolerance Planta 1997201293ndash7
Ronde JAD Cress WA Krugerd GHJ Strasserd RJ Van Staden J Photosynthetic responseof transgenic soybean plants containing an Arabidopsis P5CR gene during heat anddrought stress J Plant Physiol 20041611211ndash24
Salvi S Tuberosa R To clone or not to clone plant QTLs present and future challengesTrends Plant Sci 200510297ndash304
Sanchez AC Subudhi PK Rosenow DT Nguyen HT Mapping QTLs associated with droughtresistance in sorghum (Sorghum bicolor L Moench) Plant Mol Biol 200248713ndash26
Saranga Y Menz M Jiang CX Wright RJ Yakir D Paterson AH Genomic dissection of genotypetimesenvironment interactions conferring adaptation of cotton to aridconditions Genome Res 2001111988ndash95
Sari-Gorla M Krajewski P Di Fonzo N Villa M Frova C Genetic analysis of droughttolerance in maize by molecular markers II Plant height and 1047298owering Theor ApplGenet 199999289ndash95
Seki M Narusaka M Ishida J Nanjo T Fujita M Oono Y et al Monitoring the expressionpro1047297les of 7000 Arabidopsis genes under drought cold and high-salinity stressesusing a full-length cDNA microarray Plant J 200231279ndash92
Serraj R Sinclair TR Osmolyte accumulation can it really increase crop yield underdrought conditions Plant Cell Environ 200225333ndash41
Serraj R Krishnamurthy L Kashiwagi J Kumar J Chandra S Crouch JH Variation in roottraits of chickpea (Cicer arietinum L) grown under terminal drought Field CropsRes 200488115ndash27
Serraj R Hash CT Rizvi MHS Sharma A Yadav RS Bidinger FR Recent advances in marker-assisted selection for drought tolerance in pearl millet Plant Prod Sci 20058(3)334ndash7
Shen YG Du BX Zhang WK Zhang JS Chen SY AhCMO regulated by stresses in Atriplexhortensis can improve drought tolerance in transgenic tobacco Theor Appl Genet2002105815ndash21
Shinozaki K Yamaguchi-Shinozaki K Molecular responses to dehydration and lowtemperature differences and cross-talk between two stress signaling pathwaysCurr Opin Plant Biol 20003217ndash23
Singh KB Omar M Saxena MC Johansen C Registration of FLIP 87-59C a drought-tolerant chickpea germplasm line Crop Sci 199636(2)1ndash2
Singh SP Teran H Gutierrez JA Registration of SEA 5 and SEA 13 drought tolerant drybean germplasm Crop Sci 200141276ndash7
Singh SP Teran H Lema M Schwartz HF Miklas PN Registration of white moldresistant dry bean germplasm line A 195 J Plant Registr 2007162ndash3
Sivamani E Bahieldin A Wraith JM Improved biomass productivity and water useef 1047297ciency under water de1047297cit conditions in transgenic wheat constitutivelyexpressing the barley HVA1 gene Plant Sci 20001551ndash9
Skovmand B Reynolds MP DeLacy IH Searching genetic resources for physiologicaltraits with potential for increasing yield In Reynolds MP Ortiz-Monasterio IMcNab A editors Application of Physiology in Wheat Breeding 2001 p 17ndash28
Steele K Novel upland rice variety bred using marker-assisted selection and client-oriented breeding released in Jharkhand India Bangor University 2009
Steele KA Price AH Shashidar HE Witcombe JR Marker-assistedselection to introgressrice QTLs controlling root traitsinto an Indianupland rice variety Theor Appl Genet2006112208ndash21
Steele KA Virk DS Kumar R Prasad SC Witcombe JR Field evaluation of upland ricelines selected for QTLs controlling root traits Field Crops Res 2007101180 ndash6
Sunkar R Kapoor A Zhu JK Post transcriptional induction of two CuZn superoxidedismutase genes in Arabidopsis is mediated by down regulation of miR398 andimportant for oxidative stress tolerance Plant Cell 2006182051ndash65
Talame V Sanguineti MC Chiapparino E Bahri H Ben Salem M Forster BP et alIdenti1047297cation of Hordeum spontaneum QTL alleles improving 1047297eld performance of barley grown under rainfed conditions Ann Appl Bot 2004144309ndash20
Tanner NKCordinO BanroquesJ DoereM Linder PThe Q Motif a newlyidenti1047297ed motif in DEAD box helicases may regulate ATP binding and hydrolysis Mol Cell 200311127ndash38
Taramino G Tarchini R Ferrario S Lee M Pe ME Characterization and mapping of simplesequence repeats (SSRs) in Sorghum bicolor Theor Appl Genet 19979566ndash72
Teulat B Monneveux P Wery J Borriegraves C Souyris I Charrier A et al Relationshipsbetween relative water content andgrowth parameters in barleya QTLstudy NewPhytol 199713799-107
Thi Lang N Chi Buu B Fine mapping for drought tolerance in rice ( Oryza sativa L)Omonrice 2008169-15Till BJ Comai L Henikoff S Tillering and ecotillering for crop improvement In
Varshney RK Tuberosa R editors Genomics-assisted crop improvement genomicsapproaches and platforms vol 1 2007 p 333ndash50
Tondelli A Francia E Barabaschi D Aprile A Skinner JS Stockinger EJ et al Mappingregulatory genes as candidates for cold and drought stress tolerance in barleyTheor Appl Genet 2006112445ndash54
Tuberosa R Salvi S Genomics approaches to improve drought tolerance in cropsTrends Plant Sci 200611405ndash12
Valkoun JJ Wheat pre-breeding using wild progenitors Euphytica 200111917ndash23Vashisht AA Tuteja N Stress responsive DEAD-box helicases a new pathway to
engineer plant stress tolerance J Photochem Photobiol B Biol 200684150 ndash60Vendruscolo ECG Schuster I Pileggi M Scapim CA Molinari HBC Marur CJ et al Stress-
induced synthesis of proline confers tolerance to water de1047297cit in transgenic wheat J Plant Physiol 20071641367ndash76
Vienne D Leonardi A Damerval C Zivy M Genetics of proteome variation for QTL characterization application to drought stress responses in maize J Exp Bot 199950303ndash9
Villareal RL Mujeeb-Kazi A Rajaram S Toro ED Morphological variability in somesynthetic hexaploid wheats derived from Triticum turgidum times T tauschii J GenetBreed 1994487-16
Vinh NT Paterson AH Genome mapping and its implication for stress resistance inplants In Ashraf M Harris PJC editors Abiotic stresses plant resistance throughbreeding and molecular approaches 2005
Vinocur B Altman A Recent advances in engineering plant tolerance to abiotic stressachievements and limitations Curr Opin Biotechnol 200516123ndash32
Wang YJ Hao YJ Zhang ZG Chen T Zhang JS Chen SY Isolation of trehalose-6-phosphate phosphatase gene from tobacco and its functional analysis in yeast cells
J Plant Physiol 2005162215ndash23Wu R Garg A Engineering rice plants with trehalose-producing genes improves
tolerance to drought salt and low temperature ISB News Report 2003Xiao B Huang Y Tang N Xiong L Overexpression of LEA gene in rice improves drought
resistance under 1047297eld conditions Theor Appl Genet 200711535ndash46Xinglai P Sangang X Qiannying P Yinhong S Registration of lsquo Jinmai 50rsquo wheat Crop Sci
200646983ndash5Xiong L Lee H Ishitani M Zhu JK Regulation of osmotic stress-responsive gene
expression by the LOS6ABA1 locus in Arabidopsis J Biol Chem 20022778588ndash96Xu GW Magill CW Shertz KF Hart GE A RFLP linkage map of Sorghum bicolor (L)
Moench Theor Appl Genet 199489139ndash45Xu D Duan X Wang B Hong BHo THD Wu RExpression of a late embryogenesis abundant
protein gene HVA1 from barley confers tolerance to water de1047297cit and salt stress intransgenic rice Plant Physiol 1996110249ndash57
Yadav RS Hash CT Bidinger FR Devos KM Howarth CJ Genomic regions associatedwith grain yield and aspects of post-1047298owering drought tolerance in pearl milletacross stress environments and testers background Euphytica 2004136265ndash77
YamadaM MorishitaH Urano K Shiozaki N Yamaguchi-Shinozaki K ShinozakiK et alEffects of free proline accumulation in petunias under drought stress J Exp Bot2005561975ndash81
Yang WJ Rich PJ Axtell JD Wood KV Bonham CC Ejeta G et al Genotypic variation forglycinebetaine in sorghum Crop Sci 200343162ndash9
Yan-Ying QU Ping MU Xue-Qin L Yu-Xiu T Feng W Hong-Liang Z et al QTL mappingand correlations between leaf water potential and drought resistance in rice underupland and lowland environments Acta Agron Sin 200834(2)198ndash206
Ye XDWu XLZhaoH Frehner MNoumlsberger J Potrykus Iet al Altered fructan accumulationin transgenic Lolium multi 1047298orum plants expressing a Bacillus subtilis sacB gene Plant Cell
Rep 200120205ndash
12Zaharieva M Gaulin E Havaux M Acevedo E Monneveux P Drought and heat responses in
the wild wheat relative Aegilops geniculata Roth potential interest for wheatimprovement Crop Sci 2001411321ndash9
Zhang J Zheng HGAartiA Pantuwan G NguyenTT Tripathy JNet alLocatinggenomicregions associated with components of drought resistance in rice comparativemapping within and across species Theor Appl Genet 200110319ndash29
Zhang GH Su Q An LJ Wu S Characterization and expression of a vacuolar Na+H+
antiporter gene from the monocot halophyte Aeluropus littoralis Plant PhysiolBiochem 200846117ndash26
Zhao J QTLs for oil content and their relationships to other agronomic traits in anEuropeantimesChinese oilseed rape population Germany Diss Grorg-Agust Univer-sity of Goettingen 2002
Zhao J Ren W Zhi D Wang L Xia G Arabidopsis DREB1ACBF3 bestowed transgenic tallfescue increased tolerance to drought stress Plant Cell Rep 2007261521ndash8
Zhou W Li Y Zhao BC Ge RC Shen YZ Wang G Huang ZJ Over-expression of TaSTRG gene improves salt and drought tolerance in rice J Plant Physiol 200915(166)1660ndash70
Pratt RC Casey MA Registration of maize germplasm line Oh605 Crop Sci 2006461004ndash5Qin F Kakimoto M Sakuma Y Maruyama K Osakabe Y Tran LSP et al Regulation and
functional analysis of ZmDREB2A in response to drought and heat stress in Zea mays LPlant J 20075054ndash69
Quan R Shang M Zhang H Zhao Y Zhang J Engineering of enhanced glycinebetainesynthesis improves drought tolerance in maize Plant Biotechnol J 20042477ndash86
Quarrie SA Gulli M Calestani C Steed A Marmiroli N Location of a gene regulatingdrought-induced abscisic acid production on the long arm of chromosome 5A of wheat Theor Appl Genet 199489794ndash800
QuickJS Stromberger JA Clayshulte S Clifford B Johnson JJ Peairs FB et al Registrationof lsquoPrairie Redrsquo wheat Crop Sci 2001411362ndash3
RajaramS Prospects andpromise ofwheatbreedingin the21stcentury Euphytica 20011193-15Rajaram S Role of conventional plant breeding and biotechnology in future wheat
production Turk J Agric Forest 200529105ndash11Rami JF Dufour P Trouche G Fliedel G Mestres C Davrieux F et al Quantitative trait
loci for grain quality productivity morphological and agronomical traits insorghum (Sorghum bicolor L Moench) Theor Appl Genet 199897605ndash16
Rassaa N Salahb HBH Latiri K Thermal responses of durum wheat Triticum durum toearly water stress consequence on leaf and 1047298ower development Plant Biol Pathol2008331(5)363ndash71
Reddy LJ Nigam SN Rao RCN Reddy NS Registration of ICGV 87354 peanut germplasmwith drought tolerance and rust resistance Crop Sci 200141274ndash5
Ribaut JM Ragot M Marker-assisted selection to improve drought adaptation in maize thebackcross approach perspectives limitations and alternatives J Exp Bot 200658351ndash60
Robin S Pathan MS Courtois B La1047297tte R Carandang S Lanceras S et al Mappingosmotic adjustment in an advanced back-cross inbred population of rice Theor
Appl Genet 2003107(7)1288ndash96Romero C Belles JM Vaya JL Serrano R Culianez-Macia FA Expression of the yeast
trehalose-6-phosphate synthase gene in transgenic tobacco plants pleiotropicphenotypes include drought tolerance Planta 1997201293ndash7
Ronde JAD Cress WA Krugerd GHJ Strasserd RJ Van Staden J Photosynthetic responseof transgenic soybean plants containing an Arabidopsis P5CR gene during heat anddrought stress J Plant Physiol 20041611211ndash24
Salvi S Tuberosa R To clone or not to clone plant QTLs present and future challengesTrends Plant Sci 200510297ndash304
Sanchez AC Subudhi PK Rosenow DT Nguyen HT Mapping QTLs associated with droughtresistance in sorghum (Sorghum bicolor L Moench) Plant Mol Biol 200248713ndash26
Saranga Y Menz M Jiang CX Wright RJ Yakir D Paterson AH Genomic dissection of genotypetimesenvironment interactions conferring adaptation of cotton to aridconditions Genome Res 2001111988ndash95
Sari-Gorla M Krajewski P Di Fonzo N Villa M Frova C Genetic analysis of droughttolerance in maize by molecular markers II Plant height and 1047298owering Theor ApplGenet 199999289ndash95
Seki M Narusaka M Ishida J Nanjo T Fujita M Oono Y et al Monitoring the expressionpro1047297les of 7000 Arabidopsis genes under drought cold and high-salinity stressesusing a full-length cDNA microarray Plant J 200231279ndash92
Serraj R Sinclair TR Osmolyte accumulation can it really increase crop yield underdrought conditions Plant Cell Environ 200225333ndash41
Serraj R Krishnamurthy L Kashiwagi J Kumar J Chandra S Crouch JH Variation in roottraits of chickpea (Cicer arietinum L) grown under terminal drought Field CropsRes 200488115ndash27
Serraj R Hash CT Rizvi MHS Sharma A Yadav RS Bidinger FR Recent advances in marker-assisted selection for drought tolerance in pearl millet Plant Prod Sci 20058(3)334ndash7
Shen YG Du BX Zhang WK Zhang JS Chen SY AhCMO regulated by stresses in Atriplexhortensis can improve drought tolerance in transgenic tobacco Theor Appl Genet2002105815ndash21
Shinozaki K Yamaguchi-Shinozaki K Molecular responses to dehydration and lowtemperature differences and cross-talk between two stress signaling pathwaysCurr Opin Plant Biol 20003217ndash23
Singh KB Omar M Saxena MC Johansen C Registration of FLIP 87-59C a drought-tolerant chickpea germplasm line Crop Sci 199636(2)1ndash2
Singh SP Teran H Gutierrez JA Registration of SEA 5 and SEA 13 drought tolerant drybean germplasm Crop Sci 200141276ndash7
Singh SP Teran H Lema M Schwartz HF Miklas PN Registration of white moldresistant dry bean germplasm line A 195 J Plant Registr 2007162ndash3
Sivamani E Bahieldin A Wraith JM Improved biomass productivity and water useef 1047297ciency under water de1047297cit conditions in transgenic wheat constitutivelyexpressing the barley HVA1 gene Plant Sci 20001551ndash9
Skovmand B Reynolds MP DeLacy IH Searching genetic resources for physiologicaltraits with potential for increasing yield In Reynolds MP Ortiz-Monasterio IMcNab A editors Application of Physiology in Wheat Breeding 2001 p 17ndash28
Steele K Novel upland rice variety bred using marker-assisted selection and client-oriented breeding released in Jharkhand India Bangor University 2009
Steele KA Price AH Shashidar HE Witcombe JR Marker-assistedselection to introgressrice QTLs controlling root traitsinto an Indianupland rice variety Theor Appl Genet2006112208ndash21
Steele KA Virk DS Kumar R Prasad SC Witcombe JR Field evaluation of upland ricelines selected for QTLs controlling root traits Field Crops Res 2007101180 ndash6
Sunkar R Kapoor A Zhu JK Post transcriptional induction of two CuZn superoxidedismutase genes in Arabidopsis is mediated by down regulation of miR398 andimportant for oxidative stress tolerance Plant Cell 2006182051ndash65
Talame V Sanguineti MC Chiapparino E Bahri H Ben Salem M Forster BP et alIdenti1047297cation of Hordeum spontaneum QTL alleles improving 1047297eld performance of barley grown under rainfed conditions Ann Appl Bot 2004144309ndash20
Tanner NKCordinO BanroquesJ DoereM Linder PThe Q Motif a newlyidenti1047297ed motif in DEAD box helicases may regulate ATP binding and hydrolysis Mol Cell 200311127ndash38
Taramino G Tarchini R Ferrario S Lee M Pe ME Characterization and mapping of simplesequence repeats (SSRs) in Sorghum bicolor Theor Appl Genet 19979566ndash72
Teulat B Monneveux P Wery J Borriegraves C Souyris I Charrier A et al Relationshipsbetween relative water content andgrowth parameters in barleya QTLstudy NewPhytol 199713799-107
Thi Lang N Chi Buu B Fine mapping for drought tolerance in rice ( Oryza sativa L)Omonrice 2008169-15Till BJ Comai L Henikoff S Tillering and ecotillering for crop improvement In
Varshney RK Tuberosa R editors Genomics-assisted crop improvement genomicsapproaches and platforms vol 1 2007 p 333ndash50
Tondelli A Francia E Barabaschi D Aprile A Skinner JS Stockinger EJ et al Mappingregulatory genes as candidates for cold and drought stress tolerance in barleyTheor Appl Genet 2006112445ndash54
Tuberosa R Salvi S Genomics approaches to improve drought tolerance in cropsTrends Plant Sci 200611405ndash12
Valkoun JJ Wheat pre-breeding using wild progenitors Euphytica 200111917ndash23Vashisht AA Tuteja N Stress responsive DEAD-box helicases a new pathway to
engineer plant stress tolerance J Photochem Photobiol B Biol 200684150 ndash60Vendruscolo ECG Schuster I Pileggi M Scapim CA Molinari HBC Marur CJ et al Stress-
induced synthesis of proline confers tolerance to water de1047297cit in transgenic wheat J Plant Physiol 20071641367ndash76
Vienne D Leonardi A Damerval C Zivy M Genetics of proteome variation for QTL characterization application to drought stress responses in maize J Exp Bot 199950303ndash9
Villareal RL Mujeeb-Kazi A Rajaram S Toro ED Morphological variability in somesynthetic hexaploid wheats derived from Triticum turgidum times T tauschii J GenetBreed 1994487-16
Vinh NT Paterson AH Genome mapping and its implication for stress resistance inplants In Ashraf M Harris PJC editors Abiotic stresses plant resistance throughbreeding and molecular approaches 2005
Vinocur B Altman A Recent advances in engineering plant tolerance to abiotic stressachievements and limitations Curr Opin Biotechnol 200516123ndash32
Wang YJ Hao YJ Zhang ZG Chen T Zhang JS Chen SY Isolation of trehalose-6-phosphate phosphatase gene from tobacco and its functional analysis in yeast cells
J Plant Physiol 2005162215ndash23Wu R Garg A Engineering rice plants with trehalose-producing genes improves
tolerance to drought salt and low temperature ISB News Report 2003Xiao B Huang Y Tang N Xiong L Overexpression of LEA gene in rice improves drought
resistance under 1047297eld conditions Theor Appl Genet 200711535ndash46Xinglai P Sangang X Qiannying P Yinhong S Registration of lsquo Jinmai 50rsquo wheat Crop Sci
200646983ndash5Xiong L Lee H Ishitani M Zhu JK Regulation of osmotic stress-responsive gene
expression by the LOS6ABA1 locus in Arabidopsis J Biol Chem 20022778588ndash96Xu GW Magill CW Shertz KF Hart GE A RFLP linkage map of Sorghum bicolor (L)
Moench Theor Appl Genet 199489139ndash45Xu D Duan X Wang B Hong BHo THD Wu RExpression of a late embryogenesis abundant
protein gene HVA1 from barley confers tolerance to water de1047297cit and salt stress intransgenic rice Plant Physiol 1996110249ndash57
Yadav RS Hash CT Bidinger FR Devos KM Howarth CJ Genomic regions associatedwith grain yield and aspects of post-1047298owering drought tolerance in pearl milletacross stress environments and testers background Euphytica 2004136265ndash77
YamadaM MorishitaH Urano K Shiozaki N Yamaguchi-Shinozaki K ShinozakiK et alEffects of free proline accumulation in petunias under drought stress J Exp Bot2005561975ndash81
Yang WJ Rich PJ Axtell JD Wood KV Bonham CC Ejeta G et al Genotypic variation forglycinebetaine in sorghum Crop Sci 200343162ndash9
Yan-Ying QU Ping MU Xue-Qin L Yu-Xiu T Feng W Hong-Liang Z et al QTL mappingand correlations between leaf water potential and drought resistance in rice underupland and lowland environments Acta Agron Sin 200834(2)198ndash206
Ye XDWu XLZhaoH Frehner MNoumlsberger J Potrykus Iet al Altered fructan accumulationin transgenic Lolium multi 1047298orum plants expressing a Bacillus subtilis sacB gene Plant Cell
Rep 200120205ndash
12Zaharieva M Gaulin E Havaux M Acevedo E Monneveux P Drought and heat responses in
the wild wheat relative Aegilops geniculata Roth potential interest for wheatimprovement Crop Sci 2001411321ndash9
Zhang J Zheng HGAartiA Pantuwan G NguyenTT Tripathy JNet alLocatinggenomicregions associated with components of drought resistance in rice comparativemapping within and across species Theor Appl Genet 200110319ndash29
Zhang GH Su Q An LJ Wu S Characterization and expression of a vacuolar Na+H+
antiporter gene from the monocot halophyte Aeluropus littoralis Plant PhysiolBiochem 200846117ndash26
Zhao J QTLs for oil content and their relationships to other agronomic traits in anEuropeantimesChinese oilseed rape population Germany Diss Grorg-Agust Univer-sity of Goettingen 2002
Zhao J Ren W Zhi D Wang L Xia G Arabidopsis DREB1ACBF3 bestowed transgenic tallfescue increased tolerance to drought stress Plant Cell Rep 2007261521ndash8
Zhou W Li Y Zhao BC Ge RC Shen YZ Wang G Huang ZJ Over-expression of TaSTRG gene improves salt and drought tolerance in rice J Plant Physiol 200915(166)1660ndash70