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Chemical and Functional Properties of Food Saccharides

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© 2004 by CRC Press LLC<br />

The branch-chain lengths <strong>of</strong> phytoglycogen display a single-modal chain-length<br />

distribution, which is different from the bimodal distribution <strong>of</strong> amylopectin.<br />

7.3 ORGANIZATION OF STARCH GRANULES<br />

Starch is synthesized in granules inside amyloplasts. Starch granules isolated from<br />

different botanical sources <strong>and</strong> different organs display different shapes <strong>and</strong> sizes. 55<br />

Diameters <strong>of</strong> starch granules vary from submicrons to more than 100 µm. Examples<br />

<strong>of</strong> diameters <strong>of</strong> starch granules are potato starch 15 to 75 µm, wheat A-granules 18<br />

to 33 µm <strong>and</strong> B-granules 2 to 5 µm, maize starch 5 to 20 µm, rice starch 3 to 8 µm,<br />

<strong>and</strong> amaranth starch 0.5 to 2 µm. 55 Starch granules display spherical, oval, disk,<br />

polygonal, elongated, kidney, <strong>and</strong> lobe shapes. Leaf starch has flat-shaped small<br />

granules that are submicron in diameter. 4 Wheat, barley, rye, <strong>and</strong> triticale starches<br />

display bimodal granule size distributions: the disk-shaped, large A-granules <strong>and</strong><br />

the spherical, small B-granules. 55 Rice <strong>and</strong> oats starches are known as compound<br />

starches, which are defined as multiple granules synthesized within a single amyloplast.<br />

55 Thus, starch granules are tightly packed together <strong>and</strong> develop into polygonal<br />

irregular shapes.<br />

When viewed under a polarized light-microscope, starch granules display birefringence,<br />

known as the Maltese cross. The Maltese cross birefringence reflects the<br />

radial arrangement <strong>of</strong> starch molecules in the starch granule. The center <strong>of</strong> the<br />

Maltese cross, the hilum, is the organic center <strong>of</strong> the granule where biosynthesis <strong>of</strong><br />

the starch granule initiates. The hilum is not necessarily at the geometric center <strong>of</strong><br />

the granule; some can be located close to the end <strong>of</strong> the granule, reflecting the<br />

initiating site <strong>of</strong> the starch biosynthesis.<br />

Many starch varieties, particularly those having the A-type x-ray pattern <strong>and</strong><br />

short average branch-chain length (e.g., sorghum <strong>and</strong> maize), display pinholes on<br />

the surface <strong>of</strong> starch granules. The pinholes are open ends <strong>of</strong> serpentine-like channels<br />

penetrating into the granule 56 <strong>and</strong> are attributed to enzyme hydrolysis <strong>of</strong> starch<br />

granules. Scanning electron micrographs <strong>of</strong> cracked corn kernels shows enzymedegraded<br />

starch granules located around germs, indicating that enzyme hydrolysis<br />

<strong>of</strong> starch initiates at the location surrounding the germ to generate energy for<br />

germination. 57<br />

Surface gelatinization <strong>of</strong> starch granules by saturated neutral salt solutions, such<br />

as LiCl <strong>and</strong> CaCl 2, enables separation <strong>of</strong> starch molecules by layers from the<br />

periphery <strong>of</strong> the granule. Therefore, one can investigate structures <strong>of</strong> starch molecules<br />

at different radial locations <strong>of</strong> a granule. 58,59 The amylose content is higher at<br />

the periphery <strong>of</strong> the granule than at the hilum. This difference in the amylose content<br />

is consistent with a known fact that amylose content <strong>of</strong> starch increases with size<br />

<strong>of</strong> starch granules. The same studies also reveal that amylopectin molecules at the<br />

hilum have longer branch chains than do those at the periphery. 58,59<br />

Structures <strong>of</strong> amylopectin molecules isolated from disk-shaped, large A-granules<br />

<strong>and</strong> from spherical, small B-granules <strong>of</strong> barley, wheat, <strong>and</strong> other starches have been<br />

analyzed. Large disk-shaped granules consist <strong>of</strong> amylopectin that has fewer short<br />

branch chains than does that <strong>of</strong> small spherical granules. 57 Differences in branch<br />

structures determine the shape <strong>of</strong> amylopectin molecules, which, in turn, governs

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