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Starch Structure
Starch is the primary source of stored energy in cereal grains.
Although the amount of starch contained in grains varies, it is gener- In This Chapter:
ally between 60 and 75% of the weight of the grain and provides Basic Carbohydrate
70–80% of the calories consumed by humans worldwide. Chemistry
In addition to their nutritive value, starches and modified starches
Starch Polymer
can be used to affect the physical properties of many foods. For ex- Biosynthesis
ample, commercial starches obtained from corn, wheat, various rices,
and tubers such as potato, sweet potato, and cassava (tapioca starch) Properties of Amylose
can be used in gelling, thickening, adhesion, moisture-retention, sta- and Amylopectin
bilizing, texturizing, and antistaling applications. Starch and prod- Amylopectin
ucts derived from starch are also important in the paper and textile
industries. The unique chemical and physical characteristics of starch Starch Granules
Internal Structure of the
set it apart from all other carbohydrates.
Starch Granule
Minor Constituents of
the Starch Granule
Basic Carbohydrate Chemistry
Regardless of the botanical source, starch is basically polymers of
the six-carbon sugar D-glucose, often referred to as the “building
block” of starch. The structure of the monosaccharide D-glucose can
be depicted in either an open-chain or a ring form (Fig. 1-1). The ring
configuration is referred to as a pyranose, i.e., D-glucopyranose. The
pyranose ring is the most thermodynamically stable and is the con-
figuration of the sugar in solution. The highly reactive aldehyde
D-Glucopyranose —The ring
group at carbon number 1 (C1) on D-glucose makes it a reducing sugar. form of the monosaccharide
In biological systems, D-glucopyranose is usually present in relatively D-glucose.
small amounts compared with the levels of various disaccharides and
polysaccharides that are present. Reducing sugar —A mono-
Starch consists primarily of D-glucopyranose polymers linked to- saccharide, disaccharide,
gether by α-1,4 and α-1,6 glycosidic bonds (Fig. 1-2). In forming these oligosaccharide, or related
bonds, carbon number 1 (C1) on a D-glucopyranose molecule reacts product capable of reducing an
with carbon number 4 (C4) or carbon number 6 (C6) from an adja- oxidizing ion. A common test
for the measurement of reduc-
cent D-glucopyranose molecule. Because the aldehyde group on one
ing sugars involves the reduc-
end of a starch polymer is always free, starch polymers always have tion of cupric ions (Cu+2) to
one reducing end. The other end of the polymer is called the nonre- cuprous ions (Cu+).
ducing end. Depending on the number of polymeric branches pre-
sent in a starch molecule, there could be a large number of nonre- Glycosidic bond —Covalent
ducing ends. The glycosidic linkages in starch are in the alpha (α) linkage formed between
configuration. Formation of an α linkage is determined by the orien- D-glucopyranose units.


D -glucopyranose Depending upon the plant. fruits.4-linked D-glucopyranose molecules. Starch polymers contain only α linkages.6 linkages. susceptible to amylase enzymes.g. 6 C starch is sometimes com- H C O H H O OH pared to cellulose. branched. Amylose is an essentially linear polymer. Although a significant The α -1. whereas the amylopectin molecule is much larger and is branched. for germinating seeds and leaf tissue development) and is Amylopectin —A very large. physicochemical figuration. Amylase —Any one of several Glucose polymerization in starch results in two types of polymers. Starch functions mainly as a carbohydrate source for the growing plant (e. lyzed by amylase enzymes.4 and α -1. whereas starch polymers are usually helical. starch can be found in a variety of tissues. cellulose cannot be digested by most animals. 2 H C OH H The α linkage allows some HOCH H 1 starch polymers to form 4 C helical structures.2 / CHAPTER ONE HC = O tation of the hydroxyl 6 CH2OH 1 (–OH) group on C1 of the HCOH 5 pyranose ring (Fig.. Since the β-1. continues to represent a major area of ongoing research. This seemingly trivial difference OH H HO C C HO C C results in large differences between starch and cellu- H OH H OH lose polymers. Starch polymers can be hydro- earth.4 bonds of cellulose are not D-glucopyranose units. amylose and amylopectin. tubers. including leaves. a glucose H H polymer with β-1. and susceptibil- ity to certain enzymes. ribbon-like structure. A small number of α-1. found only in plants. most no- Fig.4-linked enzymes (see Chapter 4). 1-1). polymer of starch composed of α-1. cellulose forms a sheeted. The α configu- Cellulose —The most abundant ration and the helical geometry of starch contribute to its unique carbohydrate polymer on properties and enzyme digestibility. properties. and seeds.ration. consequently the primary source of stored energy in the plant. and microorganisms. starch-degrading enzymes common to animals. it is not completely understood and branch point. polymer of starch containing both α -1. tably in structural configu- The ring form is referred to as D -glucopyranose and can be in either the α or β con. A detailed . Because of its β configuration. often referred to as the “starch-splitting” comprising β-1. The structural differences between these two polymers contribute to Amylose —An essentially linear significant differences in starch properties and functionality. The im- 3 OH H C HCOH portance of the helical ge- O ometry of starch polymers 4 HO C C 3 2 is discussed at various HCOH 5 H OH times throughout this text. To illustrate the signifi- CH2OH CH2OH CH2OH cance of the α linkage.4 bonds C OH H C α or C OH H C β between subunits.6-linked branches may be Starch Polymer Biosynthesis present. Open-chain and pyranose ring structures of the hexose sugar D -glucose.6 linkage represents amount of information exists in the scientific literature on starch the bond at the polymeric biosynthesis in higher plants. plants. 1-1.

It is believed that these branching enzymes utilize a “cut-and-paste” mechanism that positions new “branches” onto existing α-1. STARCH STRUCTURE \ 3 description of starch 6 CH OH 2 biosynthesis in the plant is a complex process and H5 O H is beyond the scope of this book. granules.4 glucan chains onto existing α-1. H5 O H H5 O H H5 O H Starch synthesis is local- H H H ized in the chloroplasts 4 1 4 1 4 1 of green photosynthetic OH H OH H OH H tissue and/or in the O O O O 3 2 3 2 3 2 amyloplasts of non-green storage tissues.4 glucans (1). The various biosynthetic reactions responsible for starch polymer synthesis are under a great deal of metabolic control. 3 2 duced within the plas. D-glucopyranose chain. α-1. Starch synthase elongates the amylose chain by successive addition of D-glucopyranose. 4 1 α-1. . This growth includes elongation of amylose as well as for- mation of branches on the amylopectin molecule.4 and α -1. The branched polymer amylopectin is formed as branching enzymes catalyze the addition of α-1. These inher- ent differences between starches from various sources contribute to their versatility as food ingredients. starch polymers in the form of glucopyranose molecules. 4 linkages cules onto a growing Fig. onto a growing amylose chain. H OH O Reducing tids of the plant cell by a End series of complex biosyn- 6 6 CH 6 CH OH thetic pathways con. often referred to as a glu- can chain. it is easy to see why amylose and amylopectin polymers vary in size and structure depending on the plant and its metabolic requirements. dissimilarities between these two poly. 1-2. a reactive form of D-glucopyranose in the plant cell. CH2OH 2 2 trolled by key enzymes. 6 linkage OH H In general terms.6 glycosidic bonds of starch. and the exact pathway that results in polymer formation is still not completely un- derstood. Given the complexity of starch biosynthesis. α -1.4 glucans via α-1.6 linkages at the branch points. O starch polymers are pro. Properties of Amylose and Amylopectin Amyloplasts —Organelles in plant cells that synthesize Although amylose and amylopectin are both composed of D. Enzymes H OH  H OH  H OH  catalyze the addition of D-glucopyranose mole. Starch synthase is the enzyme that catalyzes the addition of adenosine-diphosphoglucose (ADP-glucose).

fats and oils.4 / CHAPTER ONE TABLE 1-1. ent on the amylose polymer (2). α -1. some alcohols.6 Molecular weight Typically <0.4 (some α-1.4-linked D-glucopyranose (Fig.” The term is usually used to describe nonpolar Amylose is considered to be an essentially linear polymer com- substances. including temperature. The formation and struc- tural integrity of amylose-lipid complexes are functions of various factors. and iodine (3). The resulting “inclusion com- Fig. The interior of the helix contains hydrogen atoms entraps a second molecular species as the “guest. Characteristics of Amylose and Amylopectin Characteristic Amylose Amylopectin Shape Essentially linear Branched Linkage α-1.and diglycerides. pH. amylose is actu- plex wherein a “host” molecule ally often helical. Simplified models for the structure of amylose are shown in Figure 1-4. Some im- portant characteristics of amylose and amylopectin are listed in Table 1-1. fatty acid components of glyc- Mono. 1-3. posed almost entirely of α-1. acterization of starch and is reviewed in greater detail in Chapter 2. has suggested that some branches are pres- for water.g. respectively. particularly mono. e. . contact and/or mixing time be- tween the host amylose polymer and the “guest” molecule.and Diglycerides — erides. a straight chain structure for the sake of simplicity. however. 1-3). Complexation with lipids. and the structure of the fatty acid or glyceride..5 million 50–500 million Films Strong Weak Gel formation Firm Non-gelling to soft Color with iodine Blue Reddish brown mers result in major differences in functional properties.” and is therefore hydrophobic.6) α-1. is a well-known property of the amylose helix.4 and α-1. Hydrophobic—“Water.4 linkages of amylose. Although typically illustrated as Clathrate—An inclusion com. that have little or no affinity Recent evidence. Glycerol molecules with one or Iodine complexation is an important diagnostic tool for the char- two fatty acids attached. AMYLOSE fearing. allowing amylose to form a type of clathrate complex with free fatty acids.

cooking and can occur quite rapidly with the linear poly- mer amylose. the hydrophobic core of the amylose helix com- under favorable conditions.. with fats and food emulsifiers such as mono. fatty acid chain of a monoglyceride. 1-5).and diglycerides can Corn starch—Common corn shift starch gelatinization temperatures. structure. The evolution of the amy- lopectin model has pro- gressed with the increasing sophistication of biochemical Fig. starches now contain as much association (i. Corn starch.) amylopectin. to a plexes with the hydrophobic constituent. than about 40% amylose. wheat starch. 1-4. and molecular dimensions of amylopectin.. gelatinized and pasted. Another well-known attribute of amylose is its ability to form a gel High-amylose corn starch — after the starch granule has been cooked. can alter the properties of the develop into more extensively ordered regions and ultimately. Starch isolated from a hybrid This property is evident in the behavior of certain amylose-containing corn plant that contains greater starches.e. . clus- ter patterns. native crystalline melt- ing. 1-5. rice starch. (Gelatiniza. retrogradation) of solubilized starch polymers after as 90% amylose. Paste —Starch in which a majority of the granules have undergone gelatinization. giving it a viscosity-forming ability. loss of birefringence. Amylose complexation crystalline order. pasting. Amylose can be depicted as either a straight chain or a to reassociate in an ordered helix. as it is often called. Some high-amylose corn sidered gelling starches. and limit retrogradation.e. branch chains. An amylose helix is complexed with the techniques. AMYLOPECTIN The literature proposes sev- eral models for helical config- urations. starch. ruption) of molecular orders within the starch granule mani- fested by irreversible changes in properties such as granular swelling. Amylose models. and particularly high. Gel formation is primarily the result of the re. mately 25% amylose and 75% tion. amylose corn starch isolated from hybrid corn plants are usually con. Two or more starch chains initially form a simple juncture point. Pasting involves granular swelling and exudation of the granular molecular components. As depicted. Retrogradation—Process dur- ing which starch chains begin Fig. STARCH STRUCTURE \ 5 Gelatinization—Collapse (dis- D. alter textural and viscosity starch composed of approxi- profiles of the resultant paste. Starch-lipid inclusion complex. and retrogradation are discussed in Chapter 3. i. which then may plex” (Fig. and starch solubilization.

either be delayed or prevented.6).. J. (7). but it results in more than 20. It has been estimated that about 4–6% of the link- ages within an average amylopectin molecule are α-1. polymer. L. namely small and large chains (5. The term “waxy” has nothing to do with the pres. For example.000. Approximate Amylose and Amylopectin Content of Common Food starch chain. Representation of a portion of an Because of the highly branched nature of amylopectin. the predominant molecule in most nor- mal starches. the MW of amylose can range from about 243. MN) sidered to be non-gelling but typically have a cohesive and gummy texture. more commonly. This unique configuration contributes to the crystalline nature of amylopectin and an ordered arrangement of amylopectin molecules within the starch granule. 1997.000–3. This may appear to be a small percentage.500–6. Although amy- Degree of polymerization lose from potato starch has been reported to have a MW of up to (DP) —The molecular size of a about 1. Starch Type (%) (%) pectin content of waxy starches Dent corn 25 75 is generally about 95% or Waxy corn <1 >99 more. R. The be- havior of branch chains of amylopectin is similar to that of amylose chains in that entire chains or. Paul. The much larger amylopectin molecule has a DP of about 300. but rather de. Tapioca 17 83 ence of wax.000 to 972. St. (Reprinted from Whistler. Pastes from starches that and BeMiller. is a branched polymer that is much larger than amylose (Fig. N. the MW of amylose is typically less than 500. Amylose from various botanical sources has a DP of about 1.000. The amylo. On the basis of these numbers and given a molecular weight (MW) for anhydroglucose of 162. Amylopectin is composed of α-1.4- linked glucose segments connected by α-1. of typical packed clusters (right). Fig. retrogradation is slowed and gel formation can ble helical. The small chains have an average degree of polymerization (DP) of about 15.. Carbohydrate Chem- istry for Food Scientists. Studies suggest a bimodal size distribution of polymer chains. whereas that of the larger chains is about 45. it refers to The MW of amylopectin can range from about 10.000.6 linkages (4). 1-6.000.000 to the number of α-1. In this case. amylopectin molecule (left) and enlargement its properties differ from those of amylose. Potato 20 80 scribes the appearance of the High-amylose corn 55–70 (or higher) 45–30 (or lower) cross section of the mutant Wheat 25 75 corn kernel from which it was first isolated. American Association contain essentially all amylopectin (waxy starches) are con- of Cereal Chemists.6 / CHAPTER ONE Amylopectin. Starches Waxy starch —“Amylopectin” Amylose Amylopectin starch from certain cereals such Content Content as corn and rice. although the branches themselves are not large. structure. 1-6). portions of chains can be helical. and pairs of chains are dou. Rice 19 81 . Individual given the size of the molecule and its “tumbleweed-like” chains are helical.6-linked branch points.000 branches in an average mol- ecule.4-linked D-glucopyranose units in a TABLE 1-2.

barley. the method of MW determination. shape. The ratio of amylose to amylopectin within a given type of starch Compound starch granules— is a very important point to consider with respect to starch function. The diameters of the granules generally range from less than 1 µm to more than 100 µm.000. but as com- ponents of discrete. Some granules. spherical. or angular) or quite ir- regular. and rye granules exhibit two separate distri- butions of granule sizes and distinctly different shapes. lenticular spherical spherical truncated. such as those in oats and rice. Approximate Size and Shape of Common Food Starch Granulesa High- Dent Waxy Amylose Property Corn Corn Corn Wheat Rice Potato Tapioca Source Cereal Cereal Cereal Cereal Cereal Tuber Root Diameter (µm) 5–30 5–30 5–30 1–45 1–3 5–100 4–35 Shape Polygonal. and and amylopectin. R. semicrystalline aggregates called starch granules. Polygonal. gelatinization and past- ing profiles. J. ovoid. Although the major components of all types of starch granules are amylose and amylopectin polymers. Round. Starch Granules Amylose and amylopectin do not exist free in nature. These are called compound starch granules. or various ratios in between. irregular compound “kettle granules drum” a Adapted from Alexander. all amylopectin. and textural attributes. Differences in the MW of amylose and amylopectin Starch granules —Naturally fractions are directly related to the plant source. The amylose and amylopectin content and structure of individual starch granules. have a higher level of structure in which many small. TABLE 1-3. the method of poly. Polygonal. more spherical granules (type B) about 3 µm in diameter can both be extracted from wheat flour. affect the architecture of the starch granule... STARCH STRUCTURE \ 7 500. round round round. For example. partially crystalline.000. The size. and structure of these granules vary substantially among botanical sources (Table 1-3). individual granules are cohesively bound together in an organized manner. 1995. Cereal Foods World 40:763–764. oval granules (type A) of approximately 35 µm (major axis) and smaller. Wheat. large.g. occurring. . Oval. it is now possible to obtain starches from various hybrid plant sources that contain essentially all amylose. The approximate amylose and amylopectin contents of several starches are shown in Table 1-2. Potato starch: New prospects for an old product. Naturally occurring aggregates ality in foods. By using classical breeding techniques as well as sophisticated molecular biology. and shapes can be regular (e. there is great diversity in the structure and characteristics of native starch granules. discrete aggregates of amylose mer isolation (usually a solvent precipitation method is used). Polygonal. Oval.

The model in Figure 1-8. if it is heated in water indefinitely. MINOR CONSTITUENTS OF THE STARCH GRANULE Proteins. These radially oriented amylopectin “clusters” are also believed to be associated with amylose. is typically less than 0. This pattern indicates that the crys- talline and amorphous areas of the granule alternate. .5% (dry basis). The location of amylose within the granule remains one of the unknown facts required to complete our picture of the internal structure of the starch granule. The values reported are not limited to the proteins and lipids associated with the granules but reflect residuals from the starch isolation (milling) process. 1-7). amylopectin is a large mol- ecule composed of two distinct populations of chain lengths. however. The smaller chains are thought to be in such close proximity that they interact strongly. Bar = 10 µm. de- broken starch granules. Moisture typically equilibrates to about 12% in a starch powder. Granules exhaustively treated in this man- ner demonstrate a ringed pattern analogous to the growth rings on a crosscut piece of wood from a tree trunk (Fig. such as α-amylase. Ash content. As discussed previously.8 / CHAPTER ONE INTERNAL STRUCTURE OF THE STARCH GRANULE The arrangement of amylose and amylopectin within the starch granule is not completely understood. has been proposed by French and Kainuma (9). resulting in crystalline regions that are quite extensive and Fig. (Reprinted scribing the arrangement of the amylopectin mole- from [8]) cule within a growth ring of a starch granule. the native granule swells until its structure eventually disintegrates. Much of what is known about the internal struc- ture is the result of microscopic evaluation of par- tially degraded granules. amorphous areas of the granule are generally degraded more easily by acid and enzymes. moisture. which is interwoven throughout the crystalline and amorphous areas. Depending on the starch. Table 1-4 summarizes the lipid and protein contents of common food starches. It is thought that this configuration results from al- ternating periods of growth and rest during the syn- thesis of the starch granule (8). lipids. although variable. This loss of internal order occurs at different temperatures for different types of starch. starch gran- ules become much less ordered. Cross section of a sorghum kernel treated arranged regularly with respect to each other with α -amylase. and amylose and amy- lopectin are released into an aqueous suspension. Compared with crystalline areas. Note the concentric rings in the throughout the granule. and ash (minerals and salts) are also present in starch granules in very small quantities. The “packaging” of these two polymers in the native starch granule is not random but is very orga- nized. When heated in the presence of water. 1-7.

4 0. for example.1 the gluten and starch. after extrac. Starch—Composition.1 Endosperm—Interior portion of the wheat kernel containing Lipid (%) 0. Although some starch sources are inherently “cleaner” with respect to their lipid and protein contents. isolated a specific protein they called the 15K starch granule protein (11). from [9]) tion of starch granule proteins in sodium dodecyl sul- fate. the most dramatic effect of these components is on the flavor profile of the starch. with permission. 1995. L. while the surface proteins are more loosely associated with the exterior of the granule. Hard wheats require more energy to mill to flour and generate more damaged TABLE 1-4.90 0. Approximate Lipid and Protein Content of Common Food Starchesa starch during the process than Dent Waxy soft wheats. Food Tech Europe. STARCH STRUCTURE \ 9 Generally speaking. the fuel source tional value in foodstuffs. “Surface” starch granule proteins can be extracted with salt solutions. the amount of residual lipid and pro- tein is a function of the efficacy of the milling (starch isolation) process. with the detergent sodium do- decyl sulfate or an alkaline solution. Compared with most cereal starches.80 0. This study involved a genetically diverse set of more than 150 wheat cultivars and strongly suggests that starch granule proteins play a significant Hardness —Amount of force required to crush a kernel of role in determining hardness. wheat. (Reprinted. tuber and root starches contain less lipid and protein compared with cereal starches. for a sprouting wheat plant.1 nm). modifications. Proteins. Because starch milling is generally a wet fractionation and pu- rification process.1 0. .1-8. comprise flour. This protein was much more abundant in starch extracted from soft wheat than in that from hard wheat. wheat endosperm. 44-52.20 0. applications and nutri. whereas “integral” starch granule proteins require more rigorous extraction. tapioca and potato starch are considered to be very bland in flavor because of the small amounts of lipid and protein present.1 0. Model of the structure of amylopectin been demonstrated to correlate well with the hardness of in starch granules (1 Å = 0. Starch granule proteins have been divided into two types on the basis of their ability to be ex- tracted from the granules (10).35 0. The presence of a specific starch granule protein has Fig. pp. It has been theo- rized that the integral proteins are embedded and possi- bly covalently bound in the amylose-amylopectin structure of the granule.25 0.. Greenwell and Schofield. in most instances the levels of these components are directly re- lated to the process used to isolate the granules. such as an alkali washing step in the milling of corn starch. Corn Corn Wheat Potato Tapioca Protein (%) 0. June/July. Novel milling techniques. Although minor amounts of residual lipid and protein can influence gelatinization. which a Adapted from Davies. can sig- nificantly reduce the levels of residual lipid and protein.

MN. There is some evidence. It is generally thought that they are highly associated with the amylose components of the starch gran- ules (12). R.-E. Ed. New York. Mercier. 6. Triglyceride—Three fatty acids Starch lipids are contained within the granules. Water and ice.. and Krisman. Ash content can vary depending upon the source of the raw ma- terial. AVI. A. 1995. The non-starch lipids have diverse structures. C. Eds. trated in the germ. however. F. J. Lineback and G. Harn. Paul. 147:342-347. R. Res. Fennema. Carbohydr. Starches also contain trace amounts of mineral elements and in- organic salts. Fennema. Robin. Most of the lipid components of cereal grains are concen- plant within the kernel. C. 4. L. these minerals and salts are referred to as ash. 2nd ed.. Wasserman. For example. Starch/Staerke 40:44-50. 1986. Cereal Foods World 40:810-817. that much of the lyso- phospholipid content is not contained within amylose inclusion complexes. tain phosphate groups and are similar in structure to the common emulsifier lecithin. Mu-Forster. E. American Association of Cereal Chemists. 7. References 1. typically calculated by deter- termed “lysophospholipids. 5. R. Starch/Staerke 47:207-209. This complex is very stable and a sample. (13) generated thermal data indicating that many of the amylose complexes do not form until the starch is gelatinized. Marcel Dekker. P. Pages 23-67 in: Food Chemistry. These lipids con- attached to a glycerol molecule. How the lysophospholipids are arranged in the starch granule is a matter of some conjecture. R. Polymodal distribution of the chain lengths of amy- lopectins. The lipids of the endosperm have been classified as starch lipids (those associated with starch granules) and non-starch Spherosomes—Fat-containing lipids (such as those contained in the spherosomes dispersed through- organelles in endosperm cells. and its significance. Current concepts of starch structure. A.10 / CHAPTER ONE Germ—Embryo of the wheat Lipids. CT. Hood. St. 2. Pages 218-224 in: Food Carbohydrates. Biotech- nology: Progress toward genetically modified starches. Curá. B. Zobel. and types of chemical modifications that may be done to the starch. Lintnerized starches: Gel filtration and enzymatic studies of insoluble residues from pro- longed acid treatment of potato starch. 8. They are Ash—Mineral and salt fraction. H. C. D. R. 1974. Cereal Chem. C. out the endosperm).” and their structure allows them to form mining the amount of residue a complex with amylose wherein the fatty acid group is aligned in the remaining after incineration of core of the amylose helix (Fig. R. 3. O. F. 1988. triglycerides and other non-polar lipids are the major components. Hoseney. Jansson. . P. dissociates or “melts” only at very high temperatures. 1994. C. 1982. Ash. tuber starches contain covalently bound phosphate. Principles of Cereal Science and Technology.5% of the dry matter.. J. agronomic practices. Charbonniere.. and Guilbot. O. 1985. Collectively. 51:389-406. Westport. Inglett. and Huang. Molecules to granules: A comprehensive starch review. The ash content of starch is typically less than 0... with one fatty acid group per molecule.. 1-5). Amylose is not strictly lin- ear. 1995. Biliaderis et al. For example.. milling procedures. Hizukuri. S. P. R.

Pages 183-247 in Starch: Chemistry and Technology. A starch granule protein associated with endosperm softness in wheat. New York. and Schofield. 12. Food Chem. Eds. and Maurice. 11. BeMiller.. 10. P. . their composi- tion and their importance for the baking properties of wheat flours.. Robinson. 1986. and Freeman. Rayas-Duarte. 1995. French. N. T. 13. P. Academic Press. 1986. J. STARCH STRUCTURE \ 11 9. 2nd ed. 72:269-274. L. The role of starch lipids among cereal lipids. R. In situ location of a starch granule protein in durum wheat endosperm by immunocytochem- istry. F. Whistler. J. F. C. L. M. and E. S. D. Biliaderis. P... J. Cereal Chem. 63:379-380. Paschall. 22:279-295. 1984. Greenwell. Cereal Chem. J. D. Acker. T. 1982. Organization of starch granules. On the multiple melting transitions of starch/monoglyceride systems.. Getreide Mehl Brot 36:291-295. C. Page.