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REVIEW

Production, structure, physicochemical and functional propertiesof maize, cassava, wheat, potato and rice starches

Jasmien Waterschoot, Sara V. Gomand, Ellen Fierens and Jan A. Delcour

Laboratory of Food Chemistry and Biochemistry, Leuven Food Science and Nutrition Research Centre (LFoRCe), KU Leuven,Leuven, Belgium

In 2012, the world production of starch was 75 million tons. Maize, cassava, wheat and potato arethe main botanical origins for starch production with only minor quantities of rice and otherstarches being produced. These starches are either used by industry as such or following someconversion. When selecting and developing starches for specific purposes, it is important toconsider the differences between starches of varying botanical origin. Here, an overview is givenof the production, structure, composition, morphology, swelling, gelatinisation, pasting andretrogradation, paste firmness and clarity and freeze–thaw stability of maize, cassava, wheat,potato and rice starches. Differences in properties are largely defined by differences in amyloseand amylopectin structures and contents, granular organisation, presence of lipids, proteins andminerals and starch granule size.

Received: October 3, 2013Revised: January 21, 2014

Accepted: January 23, 2014

Keywords:Gelatinisation / Production / Retrogradation / Starch / Structure

1 Introduction

Starch is an important source of carbohydrates in the humandiet. In addition, it is a versatile and widely used additive inthe food, paper, chemical and pharmaceutical industries.Worldwide, 75 million tons of starch were produced in 2012(http://www.zuckerforschung.at/) and marketed as native,physically or chemically modified starch but also as liquidand solid sweeteners. This paper gives an overview of theproduction, chemical composition, structure and functionalproperties of maize, cassava, wheat, potato and rice starches.

2 Starch production and uses

Of the above mentioned world starch production more thanhalf was produced in the United States. In the EuropeanUnion, 10 million tons were produced (http://www.zuck-erforschung.at/). World production is estimated to increase to

about 85 million tons by 2015. The most important botanicalorigins for producing starches are maize, cassava, wheat andpotato, respectively. Table 1 shows the estimated 2015production for each starch. Almost 80% of the starchproduction is frommaize. In the USA, mainly maize starch isproduced, although (very) small amounts of wheat, potato andrice starches are also manufactured [1]. In Europe, in additionto maize (47%) and wheat starch (39%), also potato starch(14%) and a very small amount of rice starch (<0.5%) areproduced [2] (http://www.aaf-eu.org/). Cassava starch ismainly produced in Southeast Asia and Brazil [3]. Only asmall fraction (7% for maize, 4% for cassava, 0.9% for wheatand potato and 0.007% for rice) of the raw material cropsare used for starch production.

In applications, starch is mainly used as starch derivedsweeteners and as native and modified starches. In 2011,in the European Union, 57% of the produced starch wasconverted to sweeteners, 23% was used as native starchand 20% was modified (http://www.aaf-eu.org/). Importantstarch derived sweeteners are glucose (syrups), (high)fructose (syrups), and the polyols mannitol, sorbitol andmaltitol. Maltodextrins and oligosaccharide syrups are alsoproduced [1, 4]. Native starch is used because of its thickeningand gelling capacities. However, for a number of applications,properties of native starches fail to meet process or productrequirements. This is why starches are also chemically or

Correspondence: Jasmien Waterschoot, Laboratory of FoodChemistry and Biochemistry, Leuven Food Science and NutritionResearch Centre (LFoRCe), KU Leuven, Kasteelpark Arenberg 20,B-3001 Leuven, BelgiumE-mail: [email protected]: þ32-16-32-19-97

DOI 10.1002/star.201300238Starch/Stärke 2014, 66, 1–16 1

� 2014 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim www.starch-journal.com

http://www.zuckerforschung.at/



http://www.aaf-eu.org/


physically modified. Cross-linking and substitution arecommon modifications for starches used in food production.Cross-linking of starch improves its acid, heat and shearstability, while the introduction of bulky substituents on thestarch chains reduces retrogradation [5]. The EU exportsapproximately 10% of its produced starch and starchderivatives. Of the remaining 90%, 62% is used by the foodindustry, 1% is used for feed and 37% is used by the non-foodindustry (http://www.aaf-eu.org/). Major non-food starchapplications are in the paper and board, pharmaceutical (e.g.tablet formulations, encapsulating agents), cosmetics, chem-ical (e.g. adhesives, starch-based plastics) and textile indus-tries. Modifications to meet requirements for non-foodapplications include oxidation, cationisation, copolymerisa-tion, hydrolysis and substitution [6].

3 Starch production processes

Starch production processes and their associated costsdepend on the botanical origin of the starch. Isolation ofstarch from cassava and potato tubers is relatively simpledue to their tissue structure and their relatively low proteinand fat contents [7, 8]. Isolation of cereal starches is moredifficult as higher levels of these components need to beremoved [2, 9, 10].

3.1 Maize starch

Maize starch is commercially isolated by a wet millingprocedure. First, contaminatingmaterial is removed from thebulk mass, after which the maize is steeped in watercontaining a low level of sulphur dioxide for typically 24–40 hat 48–52°C to soften the kernels and to obtain optimal millingand separation of the maize components during the wetmilling phase. During steeping, the kernels absorb water andsulphur dioxide [10]. The latter induces protein swelling anddispersion because it cleaves inter- and intramoleculardisulphide bonds and thus reduces the average MW andincreases the solubility of the proteins [11]. During steeping,lactic acid bacteria develop and produce lactic acid from theavailable sugars. This causes a drop in pH to 4–5 which is

optimal for separation of the maize protein from starch. Inaddition, lactic acid bacteria hydrolyse some highMWsolubleprotein [10]. An important side effect of the steeping processis annealing of the starch granules, which changes theirstructural properties and increases the gelatinisation temper-ature [12]. After steeping, and wet milling, the germ isremoved and starch and protein are separated from non-starch polysaccharides by sieving. After dietary fibre removal,starch is separated from protein by centrifugation orsedimentation and flash dried [10].

3.2 Cassava starch

Cassava roots are cleaned, peeled and pulverised into a pulpyslurry. Starch is then isolated at ambient temperature.The roots contain very small levels of protein (�1%) andimpurities, which can all be removed by decantation. Non-starch polysaccharides are removed by passing the slurrythrough extractors with coarse and fine screens to removeboth large and smaller molecules. The slurry is thendewatered by centrifugation and flash dried. A small levelof sulphur dioxide can be added to the process water tocontrol bacterial growth and facilitate the process [13].

3.3 Wheat starch

Wheat is dry milled to separate bran and germ from theendosperm which is recovered as flour. Different processesare used to separate starch and gluten proteins from wheatflour, i.e. dough-ball, batter, dough-batter and high pressuredisintegration processes [9]. Van Der Borght et al. [14]extensively reviewed the main processes. In these processes,flour is mixed with different amounts of water to inducegluten agglomeration or even gluten network formation. Thedough-ball and dough-batter process are carried out atambient temperature, while for the other processes warmwater (�30–50°C) is usually used [9, 14, 15]. After formationof batter or dough, starch and gluten can be separated basedon their difference in density (by centrifugation, in hydro-cyclones) or particle size (by sieving) [9, 14, 15]. The starchis then further purified with the use of hydrocyclones orseparators and decanters and dried [9].

Table 1. Production of starches of different botanical origins

Maize starch Cassava starch Wheat starch Potato starch Rice starch

Estimated world starch production 2015(million tons/year)

64.6 10.2 6.0 3.4 <0.05

World production raw material 2011(million tons/year) (www.faostat.fao.org)

880 250 704 374 723

Main production countries USA, Japan, China,South Korea [3]

Thailand, Indonesia,Brazil, China [3]

France, Germany,USA, China [3]

Netherlands, Germany,France, China [2]

Belgium [2],Thailand, Italy

2 J. Waterschoot et al. Starch/Stärke 2014, 66, 1–16



http://www.faostat.fao.org/

3.4 Potato starch

Potatoes are ground to obtain a mixture of starch granules,broken cell walls and the ‘potato juice’, a solution containingproteins, amino acids, sugars and salts. Starch granules andnon-starch polysaccharides are separated from the juice bycentrifugation. Starch and large non-starch polysaccharidesare then separated by sieving. However, some smaller non-starch polysaccharides and some proteins remain present inthe starch fraction. The remaining non-starch polysacchar-ides can be removed by centrifugation based on the densitydifference between these polysaccharides and starch. Thesoluble protein is removed in a multi-stage, countercurrentflow system. After this, water is removed with rotatingvacuum drum filters and flash drying [3].

3.5 Rice starch

Rice starch is traditionally isolated by an alkaline procedure.Broken rice, a by-product of the conversion of brown rice intowhite rice in a process referred to as milling, is steeped in a0.3–0.5% sodium hydroxide solution for 12–24 h at 20–50°C.As mentioned above, annealing of the starch granules mayoccur at the higher process temperatures. In rice, protein andstarch are strongly associated. Sodium hydroxide solubilisesthe rice protein and facilitates the isolation of starch duringsubsequent wet milling of the kernels. After wet milling,starch is kept in suspension to allow further solubilisation ofproteins. Non-starch polysaccharides are removed by filtra-tion and the slurry is washed to remove the proteins,neutralised and dried [2, 16].

4 Chemical composition of starch granules

Starch mainly consists of two polymers of a-D-glucose unitslinked by a-1,4 and a-1,6 bonds. These are the nearly linearamylose (AM) and highly branched amylopectin (AP). Inaddition, starch contains minor constituents (lipids, proteinsand minerals) of which the levels vary with the botanicalorigin (Table 2). Tuber and root starches [e.g. potato (0.1%)and cassava starch (0.2%)] usually contain less lipid thancereal starches (0.6–1.4%) [17, 18]. Lipid content is positivelycorrelated with AM content, i.e. most AM free starches

contain negligible lipid levels [18]. Wheat starch contains ahigh level of LPLs and some glycolipids, whereas lipids ofmaize and rice starches consist of FFA and LPLs [18]. Theendogenous lipids reduce swelling and leaching of carbo-hydrates during heating of starch in excess water by theformation of AM lipid complexes [19–21]. In general, protein(0.1–0.5%) and ash (0.1–0.3%) contents of starch are verylow [22–26]. Potato starch contains a relatively high level ofphosphorus (0.09%) in the form of phosphate monoestersthat are primarily covalently bound to AP [27]. The phosphateis mainly ester linked at the C-6 (61%) (see Fig. 1) and C-3(38%) positions, with only 1% linked at the C-2 position [28].The presence of phosphate monoesters in potato starch haslarge consequences for its swelling behaviour. Negativelycharged phosphate groups cause repulsion between adjacentAP chains and allow rapid hydration and large swelling of thegranule [27]. Phosphorus in cereal starches (0.01–0.07%) ismainly present in the form of phospholipids [26, 29].

5 Amylose and amylopectin

The AM content of normal starches varies between 14 and29% [30–36]. Table 3 shows AM contents of potato, cassava,wheat, maize and rice starches. The AM content of ricestarches varies from 0 to 40% [30, 37]. Variations in AMcontent can be produced through cross-breeding, mutagene-sis or transgenic breeding [38]. AM free starches are called‘waxy’ and exist for maize, cassava, wheat, potato and ricestarches [30, 31, 39–45]. Starches with a high AM content(>30%) are also available. For maize, starches with anAM content ranging from 50 to 90% are commercially

Table 2. Composition of starches of different botanical origins


Lipids (%) 0.6–0.8 [18, 26] 0.2 [17] 0.8–1.2 [18] 0.1 [17, 26] 0.6–1.4 [18]Proteins (%) 0.4 [22, 26] 0.3 [23] 0.2–0.3 [22, 24] 0.1 [22, 26] 0.1–0.5 [25]Ash (%) 0.1 [22, 26] 0.3 [23] 0.2 [22] 0.3 [22], 0.2 [26] 0.1 [25]Phosphorus (%) 0.02 [29], 0.01 [26] 0.01 [29] 0.05 [29] 0.09 [29], 0.06 [26] 0.07 [29]

Lipid, protein, ash and phosphorus contents are shown as % of total dry weight.

Figure 1. Structure of glucose-6-phosphate in an a-(1,4) boundglucose chain.

Starch/Stärke 2014, 66, 1–16 3


available [44, 46]. Also for potato starch an AM content from56 to 92% was obtained [31, 47–49], while the highest AMcontent obtained so far for wheat (74%) [50] and rice(56%) [51] is lower. Mutants with only slightly higher AMcontents than those of their regular counterparts have beendeveloped for wheat (31–56%) [41, 47, 52–54], cassava (28–36%) [39, 55] and potato (27–37%) starches [31].

In essence, AM is a linear polymer. It contains hardly anya-1,6 branch points (<1%) [56]. AP is highly branched, with5–6% a-1,6 linkages [18]. A starch characteristic is its DP, i.e.the number of glucose units in the polysaccharide. Averagedegrees of polymerisation can be calculated based on thenumber or on the weight of the molecules. The numberaverage DP (DPn) and the weight average DP (DPw) are given

by formulas (1) and (2):

DPn ¼P1

1 DPiðmi=MiÞP11 ðmi=MiÞ ð1Þ

DPw ¼P1

1 DPimiP11 mi

ð2Þ

withmi themass concentration andMi theMWof chains witha DP¼ i [57]. DPw is always higher than DPn, except for amonodisperse polymer, in which case DPn equals DPw [58].Table 3 lists DPn of AM and AP of the different starches. DPn

of AM varies from 570 to 8025 [59–65], while DPn of AP variesfrom 4700 to 18 000 [62, 63, 66–68]. DPn values of potato and

Table 3. Contents and structural properties of amylose (AM) and amylopectin (AP) and degree of crystallinity of starches of differentbotanical origins


AM content (%) 23 [34],28 [32]

18–24 [33],18 [31, 34]

25 [35],26 [32, 34]

19–22 [31],17 [34],23 [32],25–27 [35]

17–21 [32],21 [34],14–29 [30],16–19 [36],4–16 [37]

DPna) of AM 960 [61],

830 [63]2660 [59],3642 [60]

570 [59]3827 [60],1290 [61],830–1570 [62],1200–1500 [67]

4920 [59],8025 [60]

920–1110 [64],847–1118 [65]

DPn of AP 5100 [63],15 900 [66]

– 5000–9400 [62],13 000–18 000 [67]

11 200 [66] 4700–12 800 [68],8200–10 900 [66]

Number of AMmolecules per gstarch �1017

9–10 2–3 3–11 1–2 6–9

Number of APmolecules per gstarch �1017

2–6 – 2–6 3 2–4

CLb) of AM 335 [61],340 [63]

340 [59] 250–320 [67],135–255 [62],270 [61],300 [59]

670 [59] 230–370 [64]

CL of AP 28 [70],24 [34],20–21 [63],20 [72, 75]

26 [70],28 [34],18–19 [73],19 [72]

25 [70],23 [34],19–20 [67],19–21 [62],19 [72],23 [69]

34 [70],29 [34],31 [71],23 [72]

25–28 [70],23 [34],17–18 [73],19–22 [68],18–19 [72]

Average number ofchains per moleculeof AM

2.9 [61],2.4 [63]

7.8 [59] 4.4–5.2 [67],5.5–6.5 [62],4.8 [61],1.9 [59]

7.3 [59] 2.5–4.3 [64]

Average number of chainsper molecule of AP

240 [63] – 660–920(based on [67])

500 (based on[72] and [66])

220–700 [74]

Degree of crystallinityas determined withX-ray diffraction (%)

27 [86],40 [85]

24 [86],38 [85]

20 [86],36 [85]

24 [86],28 [85]

38 [85]

a)DPn, number degree of polymerisationb)CL, average chain length



cassava AM are higher than those of maize, wheat and riceAM. Based on DPn and the contents of AM and AP, thenumber of AM and AP molecules per mass unit of starch canbe calculated. It can be estimated from data in Table 3 that agiven mass of regular starch contains more AM moleculesthan AP molecules (except for potato starch), which is due totheir lower DPn.

The average chain length (CL) of AM varies from 135 to670 glucose units per chain. The CL of potato AM is 670glucose units while that of the other starches ranges from 135to 370 glucose units [59, 61–64, 67]. Because of the highlybranched nature of AP, its CL (17–34 glucose units) is muchsmaller than that of AM. Potato and cassava AP have a slightlyhigher CL than AP of the cereal starches [34, 62, 63, 67–75].AP of high AM starches has a similar or higher CL than AP ofthe regular counterparts. In addition, these starches containsomematerial with DPs and branching patterns intermediatebetween those of AM and AP [46, 47, 76–79]. The averagenumber of chains per molecule can be calculated from DPn

and CL and is much smaller for AM (1.9–7.8) than for AP(220–920) [59, 61–64, 66, 67, 72, 74].

With regard to determination of starchmolecular size, it isimportant to note that the analytical results depend on theprocedure used. Molecular size determinations requirecomplete dissolution of the starch polymers, with removalof non-starch components that interfere with the analysis andwithout starch loss or degradation during the procedure.These requirements are challenging, so when interpretingresults, caution should be taken when comparing resultsobtained by different procedures [80, 81].

6 Different levels of starch granuleorganisation

Plants synthesise starch in a granular form. Starting from thehilum, starch is deposited in alternating amorphous and semi-crystalline concentric growth rings [16]. The semi-crystallinegrowth rings consist of amorphous and crystalline lamellae.AP is largely responsible for the crystalline character of starch.Side chains of AP form double helices which are ordered inclusters. The crystalline lamellae contain the double helices,while the amorphous lamellae contain the AP branch points,which connect the double helices [82]. The amorphous growthrings consist of AM and less ordered AP [16]. AM and AP arenot present in separate regions, but highly intermingled inthe granule [83, 84]. The degree of crystallinity is usuallydetermined with X-ray diffraction [18]. It varies from 20 to40% depending on the botanical origin (Table 3) [85, 86].

The packing of AP double helices can give rise to differentcrystal structures or polymorphic forms. Cereal starchcrystals are generally packed according to the A-type packing.Such packing is more dense than the B-type packing of e.g.potato starch and high AM maize starches. Cassava starch

contains A-type or C-type crystals (a mixture of A- and B-typecrystals) [85, 87, 88]. In A-type starches, crystals are packed ina monoclinic unit cell (a¼ 2.124 nm, b¼ 1.172 nm, c¼ 1.069nm and g¼ 123.5°) with eight water molecules, whereas inB-type starches, crystals are packed in a hexagonal unit cell(a¼ b¼ 1.85 nm and c¼ 1.04 nm) with 36 water molecules(see Fig. 2). Besides A- and B-type crystals, a third polymorphexists, i.e. V-type crystals. In this polymorph, AM singlehelices form inclusion complexes with e.g. iodine, alcoholsor fatty acids [18].

7 Morphology of the starch granule

The size and shape of starch granules (Table 4) depend onthe botanical origin and vary widely. Figure 3 shows thegranular morphology of potato, cassava, wheat, maize andrice starches. Potato starch has very large, round or ovalgranules (10–100mm), while rice starch has very small,polygonal granules (3–8mm) [32, 89, 90]. Cassava starch hasround or truncated granules while maize starch granules arepolygonal. Both starches have granules with somewhatsimilar dimensions (5–20mm for maize starch and 3–32mmfor cassava starch) [31–33, 89]. While the shape and size ofwaxy maize starch granules resemble those of regular maizestarch granules, high AMmaize starches contain, in additionto the normal polygonal granules, a number of filamentouselongated granules [89, 91]. Wheat starch has a bimodal sizedistribution, with small, round B granules (2–10mm) andlarge, lenticular (20–32mm) A granules [32, 89, 92, 93].

Figure 2. Monoclinic unit cell of A-type crystals and hexagonal unitcell of B-type crystals. Projection of the structure is in the ab plane.Reprinted from International Journal of Biological Macromolecules,23, Buléon, A., Colonna, P., Planchot, V. and Ball, S., Starchgranules: structure and biosynthesis, 85–112, Copyright (1998),with permission from Elsevier.



Average granule diameters can be calculated based on thenumber of granules or on their volume (or weight). Numbermean diameter (D [1,0]) and volume mean diameter (D [4,3])can be calculated with formulas (3) and (4),

D 1; 0½ � ¼Pn

1 din

ð3Þ

D 4; 3½ � ¼Pn

1 d4iPn

1 d3i

ð4Þ

where di is the diameter of particle i and n is the total numberof particles. Based on the number mean diameter, thenumber of granules per gram starch can be determined. Thisallows estimating that a given weight of rice starch containsabout 3000 times more granules than a same given weight ofpotato starch. The specific surface area, i.e. the total surfacearea per unit of volume, of rice starch is about ten times thatof potato starch. The values for specific surface area in Table 4are probably an underestimation, as the presence of channelsand pores on the granule surface is not included in thecalculation. However, to the best of our knowledge, no

information is available on this. Especially cereal starchescontain pores [94, 95], while for potato and cassava starchesalso depressions and protrusions on the granule surfacehave been observed [96].

8 Gelatinisation properties

Heating starch in excess water (>1:2 starch:water) above acertain temperature, the ‘gelatinisation temperature’, dis-rupts the molecular order of the granules and melts thecrystallites [97]. When relatively less water (<1:2 starch:water)is available, gelatinisation is partly postponed to highertemperatures [16]. Table 5 lists gelatinisation onset (To), peak(Tp) and conclusion (Tc) temperatures and melting enthalpies(DH) of the different starches in excess water. DH representsthe amount of energy needed to melt all the crystals [16].Wheat starch has the lowest gelatinisation temperature,followed by potato, cassava and maize starches [22, 25, 26, 30,31, 34, 71, 98–106]. Rice starches show a high variation ingelatinisation temperature, which is at least partly due to thehigh variation of AM content in regular rice starches [25, 30,

Table 4. Morphological properties of granules of starches of different botanical origins


Shape Round, polygonal [32, 89] Round, truncated [31] Round, lenticular [32, 89] Round, oval [89] Polygonal [32, 89]Diameter, range (mm) 5–20 [89] 3–32 [33] A: 20–35 B: 2–10 [32] 10–110 [31] 3–8 [32, 89]Volume mean diameter (mm) 15 [32] 17–18 [31] A: 21–23 B: 6–7 [92, 93] 48–60 [31] 6 [90]Number mean diameter (mm) (own data) 13 11 A: 17 B: 3 29 2Number of granules per g starch �108 6 10 A: 2 B: 80 0.5 1600Specific surface area (m2/kg) 270 220 A: 180 B: 670 75 670

Figure 3. SEM pictures of maize (a), cassava (b), wheat (c), potato (d) and rice (e) starches (own data). Bars represent 10mm.



34, 37, 104]. For a single starch granule, loss of molecularorder typically takes place over a very small temperature range(<1°C) [107], while gelatinisation occurs over a wider intervalfor a population of granules. The gelatinisation temperaturerange of starches mostly varies from 8 to 12°C [22, 25, 26, 30,31, 34, 37, 71, 98–105, 108], although some exceptions with alarger temperature range have also been described [30, 31,98]. DH of potato starch is slightly higher than that of theother starches. Wheat and maize starches have a low DH [22,25, 26, 30, 31, 34, 37, 71, 98–105, 108].

The gelatinisation temperature is a measure of crystalquality, while DH is a measure of both crystal quality andquantity [109]. The presence of a relatively high amountof short AP chains (DP< 14) reduces the gelatinisationtemperature, while a relatively high amount of longer chainsleads to an increased gelatinisation temperature [30, 31, 105].According to Gidley and Bulpin [110], a chain DP of at least 10is needed to form double helices. Consequently, starches witha relatively high level of short AP chains have lower crystallineorder, which leads to a lower gelatinisation temperature.Relatively high amounts of longer chains may be responsiblefor both better stabilisation of the crystal structure over alonger distance as well as for a higher gelatinisationtemperature [31]. Although CL of potato AP (23–34) ishigher than CL of the other starches (17–28) (Table 3), itsgelatinisation temperature is relatively low (To¼ 57–66°C)(Table 5), probably because of the presence of phosphatemonoesters and the more open crystal structure of B-type

than of A-type starches [26, 34, 88]. The importance of the APchain length for gelatinisation temperature is expressed bythe Gibbs–Thomson Eq. (5) for lamellar crystallites

Tm ¼ T0m 1� 2g

DHlc

� �ð5Þ

This equation relates the melting temperature (Tm) to theaverage crystalline layer thickness (lc), the melting tempera-ture of an ideal crystal with an infinite crystal size (T0

m), thelamellar surface free energy (g) and DH. Longer AP chainslead to increased lc and thus also Tm. A high Tm is obtainedwhen branch chain lengths are relatively long (high lc) andwhen crystal quality is high (low g and high DH) [111].

Branch chain length can also influence DH. Starches withhigher CL (e.g. potato starch) appear to have a higherDH [34].DH is also correlated with crystallinity, i.e. a higher degree ofcrystallinity (e.g. waxy starches) leads to a higher DH [26, 31,34, 105]. Furthermore, the presence of lipids in cerealstarches might also explain the difference in DH betweenpotato and cereal starches. The exothermic formation of AMlipid complexes can occur simultaneously with gelatinisation,thereby lowering the measured DH [22].

9 Swelling power and solubility

At room temperature, starch granules can absorb up to 30% oftheir weight in excess water without swelling noticeably [112].

Table 5. Gelatinisation properties of starches of different botanical origins in excess water measured with DSC

Reference Starch-to-water ratio To (°C)a) TP (°C)

b) Tc (°C)c) Tc� To (°C) DH (J/g)d)

Maize starch [22, 26, 34] 1:3 64–67 68–71 72–75 8–11 11–12[98] 3:7 63 67 72 9 9[103] 1:4 67 71 – – 12[104] 1:9 66 71 – – 12

Cassava starch [31, 34] 1:3 55–64 61–68 71–74 10–16 15–19[98] 3:7 60 65 75 15 11[103] 1:4 64–66 68–70 – – 11–14[104] 1:9 65 71 – – 13

Wheat starch [22, 34, 100] 1:3 53–62 61–65 64–69 7–12 9–12[98, 105, 108] 3:7 48–57 54–62 58–68 9–11 7–15[103] 1:4 60 65 – – 10

Potato starch [22, 26, 31, 34, 71] 1:3 58–63 61–68 68–73 8–11 15–24[98, 99, 101, 102] 3:7 57–66 61–70 67–75 7–12 12–18[103] 1:4 62 65 – – 17[104] 1:9 59 64 – – 17

Rice starch [34] 1:3 70 76 80 10 13[37] 3:7 61–76 67–79 72–85 8–13 8–14[25] 3:11 51–70 58–74 64–79 9–13 8–12[104] 1:9 58 65 – – 12[30] 1:2 57–76 63–79 71–83 8–19 17–20

a)To, gelatinisation onset temperature;b)Tp, gelatinisation peak temperature,c)Tc, gelatinisation conclusion temperature;d)DH, gelatinisation enthalpy.



However, during heating, starch granules absorb muchmore water and swell. At higher temperatures, part ofthe polysaccharides go into solution and leach out of thegranules [113]. Table 6 lists the swelling powers andsolubilities of the different starches. The former are theamounts of water a starch can absorb per gram starch at acertain temperature and at a certain starch concentration,while the solubilities represent the percentages of leached AMand AP at this temperature.

Potato starch has a much higher swelling power thanother starches [103, 114–116]. As mentioned above, this islargely due to its negatively charged phosphate monoesters.The swelling power of cassava starch is also higher than thatof the cereal starches [37, 103, 105, 114, 115, 117]. Granuleswelling is mainly attributed to AP and is inhibited byAM [118]. As a result, waxy starches have a higher swellingpower than their AM containing counterparts [34, 114, 117,118]. Removal of endogenous starch lipids reduces formationof AM lipid complexes and increases the swelling power ofwheat and maize starches but not up to the level of swellingthat characterises tuber starches [21, 118]. The impact ofexogenous lipids on swelling of starch granules depends onthe type of lipid added and the temperature [19, 119–121]. Theabsence of AM lipid complexes in potato and cassava starchesalso contributes to their extensive swelling power [34, 114].

Starch solubility, determined at temperatures from 84 to95°C, varies from 3 to 37%, with values for most starchesbetween 10 and 20% (Table 6) [103, 105, 114–117, 122]. BothAM and AP leach out of granules of regular starches, whileevidently only AP leaches from waxy starches. Usually, inexcess water, AM leaching starts at relatively low temper-atures (<70°C), while AP only leaches out at highertemperatures (>90°C). This has been observed for potatoand cassava [114], rice [117], maize [123] and wheat [21, 119,123] starches.

10 Pasting properties

Pasting can be described as a term encompassing the eventsthat occur after gelatinisation in a starch suspension, i.e.further swelling of the granules, leaching of polysaccharides,increase in viscosity and formation of an AM gel network [97].Changes in viscosity depend on the concentration of thestarch suspension and can be measured with a Rapid ViscoAnalyser or a Viscoamylograph. Figure 4 shows typicalpasting profiles for maize, cassava, wheat, potato and ricestarches (own data). The pasting temperature is that at whichan onset in viscosity rise can be observed. Table 7 lists pastingtemperatures, and peak and cold paste viscosities of thedifferent starches. While the term peak viscosity speaks foritself, cold paste viscosity is that measured at the end of the

Table 6. Swelling power and solubility of starches of different botanical origins

Reference Starch concentration (%) Temperature (°C) Swelling power (g/g) Solubility (%)

Maize starch [103] 0.5 84 16 –

[115] 1.0 95 23 17[122] 2.0 95 11 18

Cassava starch [103] 0.5 84 40–60 –

[114] 0.3 90 50 13Wheat starch [103] 0.5 84 40 –

[105] 2.0 90 13–25 –

[122] 2.0 95 9 20Potato starch [103] 0.5 84 168 –

[116] – 90 26–49 3–37[114] 0.3 90 60–130 12–17[115] 1.0 95 91 12

Rice starch [117] 1.0 95 25–45 13–32[37] 2.0 90 17–39 –

Figure 4. Pasting profiles of 8.0% suspensions of potato, cassava,maize, rice andwheat starches in water measuredwith a rapid viscoanalyser. The following temperature-time profile was used at astirring speed of 160 rpm: 1min at 50°C, heating from 50 to 95°Cin 9min, 10min at 95°C, cooling from 95 to 50°C in 15min,10min at 50°C (own data).



heating and cooling cycle. Potato and cassava starches havelower pasting temperatures than cereal starches. Due to thehigh level of negatively charged phosphate groups in potatostarch, viscosity development starts at lower temperatures.Potato starch reaches a very high peak viscosity. The peakviscosities of cereal starches are rather low because of theirlower swelling power [34, 37, 105, 116, 124, 125]. Waxystarches reach higher peak viscosities than their AMcontaining counterparts due to their higher swellingpower [117]. It is of note that starches with a high swellingpower are more sensitive to granule breakdown at hightemperatures than those with a lower swelling power. Pastingprofiles of potato starch show a large breakdown during the

holding phase at 95°C [114, 124]. Waxy starches also have lowthermal stability and quickly disintegrate during pasting athigh temperatures [22].

During cooling of a starch suspension, viscosityincreases as the leached AM forms a gel network [126].Cold paste viscosities of regular wheat, maize and ricestarches are at least as high as their peak viscosities, whilecold paste viscosities of potato and cassava starchesare much lower than their peak viscosities due to significantgranule breakdown [34, 105, 116, 124, 125]. After somehours, AM forms stable crystalline structures, which areacid resistant and have a melting temperature of about150 °C [16, 126]. AM crystallisation is largely responsible for

Table 7. Pasting properties of starches of different botanical origins

Reference Starch concentration (%) Pasting temperature (°C) Peak viscosity (mPa s) Cold paste viscosity (mPa s)

Maize starch [124] 8 82 2100 2000[34] 8 82 1800 2000

Cassava starch [124] 8 67 2300 1400[34] 8 68 2100 1300

Wheat starch [34] 8 89 1250 1850[125] 9 – 2100 2950[105] 11 82–90 2250–3400 2600–4000

Potato starch [124] 8 67 9500 3400[34] 8 64 8400 2750[116] 11 65–70 4100–7200 2300–3400

Rice starch [124] 8 71 2500 2050[34] 8 80 1350 1900[37] 6 72–80 1000–2400 1500–3500

Table 8. Retrogradation properties of starches of different botanical origins measured with DSC

Reference Starch-to-water ratio Storage conditions [time (days)/temperature (°C)] DH (J/g)a)

Maize starch [34] 1:3 7/4 5.8[98] 3:7 7/�22 3.0

Cassava starch [114] 1:3 28/20 No retrogradation detected[34] 1:3 7/4 3.7[98] 3:7 7/�22 No retrogradation detected

Wheat starch [34] 1:3 7/4 3.6[98] 3:7 7/�22 2.0[105] 3:7 7/4 0.7–3.0[108] 3:7 28/5 10.1–10.6

Potato starch [114] 1:3 7/20 2.8–7.0[114] 1:3 28/20 6.3–9.9[34] 1:3 7/4 7.5[98] 3:7 7/�22 4.2[99] 3:7 14/4 6.4–8.6

Rice starch [34] 1:3 7/4 5.3[140] 1:2 28/RTb) 1.4–3.1, 7.8–11.6[140] 1:2 28/6 5.7–11.1

a)DH, melting enthalpy of retrograded amylopectin;b) RT, room temperature.



the initial firmness of a starch gel, while AP retrogradation(cf. infra) is responsible for the long-term changes in gelfirmness [127].

11 Amylopectin retrogradation

Retrogradation is the process of crystallisation of APmolecules in a starch paste [16]. The term retrogradation,which means ‘to go back’, is here only used for AP and notfor AM, as strictly speaking only AP molecules can go backto a crystalline entity [16]. AP crystals are acid labile andmelt at 50–60°C [87]. Table 8 shows the melting enthalpiesof retrograded amylopectin. The extent of retrogradationhighly depends on the storage time and temperature, starchconcentration and starch structural properties [45, 128].It consists of three steps: nucleation, propagation andmaturation. Nucleation occurs faster at a relatively lowtemperature, close to the Tg of the starch, while propagationand maturation occur faster at a temperature close to themelting temperature [129]. Besides storage temperature,also the starch-to-water ratio has an important effect onretrogradation. Water content should neither be too high(>80%) nor too low (<30%) to allow retrogradation [130].Evidently, AP structure also impacts retrogradation. ShortAP chains (DP 6–9) are less susceptible to retrogradationthan chains with DP 14–24 [114, 131]. As mentioned earlier,at least a DP of 10 is needed for forming doublehelices [110]. This could explain the higher extent ofretrogradation of potato starch than of cereal starches underthe same conditions (Table 8) [35, 132]. During storage ofstarch gels, interactions or even co-crystallisation with AMcan occur, especially when the AM content is relativelyhigh [133–136].

12 Starch functionality

12.1 Paste firmness

As mentioned above, due to AM crystallisation (�short-term)and AP retrogradation (�long-term), firmness of starch gelsincreases with time [115, 127]. This is an important aspect ofstarch functionality in food products. Gel firmness can bemeasured by compressing a gel by a certain percentage. Itdepends on starch concentration, storage time and tempera-ture [137]. Table 9 lists the firmness of gels of maize, cassava,wheat, potato and rice starches. As different starch concen-trations and storage conditions are used, it is difficult tocompare results. When comparing firmness values obtainedunder the same conditions, maize, wheat and potato starchesshow similar firmness values which are higher than that ofa cassava starch gel [138, 139]. This could be due to therelatively low AM content of cassava starch and the large lossof granular integrity in the gel [139]. Rice starch gels have abroad range of firmness values (0.09–7N) [140]. AM contentis positively correlated with gel firmness. Waxy starch gelshave only a low firmness as a result of their poor gel networkformation [140]. In contrast, lipid content is negativelycorrelated with gel firmness. The formation of AM lipidcomplexes reduces the amount of AM available for networkformation [122, 141, 142].

12.2 Paste clarity

Another important characteristic in many starch applicationsin food systems is paste clarity. The presence of relativelyshort chains of AM or AP adds to opacity in food products.While for a range of products including sauces, dressings andpuddings this is not a problem, products such as fruit fillings

Table 9. Firmness of gels of starches of different botanical origins

Reference Starch content (%) Storage conditions [time (days)/temperature (°C)] Compressed proportion of gel (%) Gel firmness (N)

Maize starch [138] 8 1/RTa) 40 0.64[138] 8 7/4 40 1.03[139] 6 1/4 33 0.93[141] 6 0.2/25 11 0.04[141] 6 1/4 11 0.05

Cassava starch [138] 8 1/RTa) 40 Not measurable[138] 8 7/4 40 0.35[139] 6 1/4 33 0.19

Wheat starch [138] 8 1/RTa) 40 0.69[138] 8 7/4 40 0.70[125] 9 1/4 Not reported 0.50

Potato starch [138] 8 1/RTa) 40 0.56[138] 8 7/4 40 0.71

Rice starch [140] 8 2/6 10 0.09–4.17[140] 8 14/6 10 0.12–7.05

a) RT, room temperature.



and jellies require the starch pastes to be of high clarity [143].Paste clarity can be determined by measuring the lighttransmittance (at 650 nm) of a 1.0% starch paste. Potatostarch (42–96% light transmittance) has the highestpaste clarity, followed by cassava starch (51–81% lighttransmittance) and the cereal starches (13–62% lighttransmittance) [144–148]. The clarity of waxy cereal starchgels is better than that of their AM containing counter-parts [144–146]. The very high paste clarity of potato starchgels is caused by the absence of granule remnants in thegel (cf. high swelling power and high granular break-down) [144]. Pastes of cassava and waxy starches alsoshow few granule remnants. However, more interactionsbetween leached material lead to a higher opacity of thesegels. The low clarity of pastes of regular cereal starchesis caused by swollen granule remnants [144] and by thepresence of AM lipid complexes [149]. During storageof starch gels, their paste clarity decreases as more andmore association of AM and/or AP molecules takesplace [146, 150].

12.3 Freeze–thaw stability

Freeze–thaw stability is an important quality aspect of starchgels. When a starch gel undergoes repeated freezing andthawing cycles, it releases water. The gel is then said toundergo syneresis. The extent of syneresis is a measure for itsfreeze–thaw stability [128, 151]. During freezing, phaseseparation takes place as ice crystals are formed. As a result,starch is concentrated in the unfrozen matrix. Duringthawing of the gel, association of AM and AP takes place inthe more concentrated zones. As a result, increasinglyinsoluble aggregates are formed. Ice crystals melt and thewater is not reabsorbed by the starch gel. A sponge likestructure develops and the melted water readily separatesfrom the gel [152].

Different factors have an influence on the freeze–thawstability. These include the freezing rate, the botanical originof the starch and its AM content, the starch concentration, thenumber of freeze–thaw cycles and the sample prepara-tion [128, 153]. The freezing rate is an important factoraffecting syneresis. Slow freezing favours formation of largeice crystals which disrupt the gel structure to a larger extentthan small ice crystals [128, 152, 154]. Slow freezing also leadsto maximal ice formation and AM self-association whichresult in significant structure loss and syneresis. The latter ispositively correlated with AM content [153]. Native, AMcontaining starches have very low freeze–thaw stability.Syneresis readings after several freeze–thaw cycles for maizestarch (47–79%) [153, 155, 156], cassava starch (50–67%) [153,155], wheat starch (44–68%) [153, 155, 156] and potato starch(60–76%) [153, 155, 157] are very high. Rice starch shows abroad range of syneresis values (7–75%) [153, 155, 158, 159].Waxy starches are intrinsically more stable than AM

containing starches, although they can also undergo strongtextural changes during freeze–thaw cycles [151, 155]. Waxyrice starch has a very good freeze–thaw stability. A relativelyhigh proportion of AP side chains with DP 6–12, which is thecase for waxy rice starch, is believed to be at the basis of areduced extent of syneresis [153, 157]. In contrast, syneresisof waxy maize starch gels is comparable to that of gels fromregular maize starch [153, 155]. For different waxy maizestarches, no correlation was observed between syneresis andretrogradation enthalpy as measured with DSC. Syneresisvalues were alreadymaximal, when little, if any, double helicalorder was present. This indicates that the early stages ofAM and/or AP association (before the formation of realcrystal structures) already cause syneresis due to networkformation [160].

The presence of lipids in cereal starches may alsocontribute to a relatively high freeze–thaw stability. Granularswelling and leaching of AM are reduced in the presence oflipids. As a result, starch molecules remain close to eachother in the granules. This facilitates their reassociation andin this way it may contribute to a low freeze–thawstability [153].

In conclusion, there are significant differences instructural, physicochemical and functional properties ofstarches of different botanical origins. Starch is widely usedby the food and non-food industry in a broad range ofproducts. The overview of starch properties provided in thisreview can be of assistance when developing starches forspecific purposes. The botanical origin of the used starch hasa great impact on final product properties. Within a botanicalorigin, in some instances the starch AM content is a majordeterminant of its properties and starch should therefore beselected with great care.

The authors gratefully acknowledge Flanders’ FOOD (Brus-sels, Belgium) and the Methusalem programme ‘Food for thefuture’ (KU Leuven) for financial support. J.A.D. is W.K. KelloggChair in Cereal Science and Nutrition at the KU Leuven.

The authors have declared no conflict of interest.

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