Degree in vitro,

J#{246}rgen Ho/rn, Ann-Charlotte

of starch gelatinization, and metabolic response

MSc; Jngmar Lundquist, MD, Eliasson, PhD; and Nils-Georg
ABSTRACI common degree were studied. of starch Glycemic ready-to-eat Wheat foods gelatinization, starch with response

digestion in rats12
Bj#{246}rck, PhD; PhD
of

rate

of starch

PhD; Inger Asp, MD,

after

is only

Starch in many the relationships among in vitro digestion rate, and in vivo metabolic response in rats different degrees ofgelatinization was used in the experiments.
ingestion starchy partly gelatinized.

foods varies.

In view ofthis,

Plasma glucose and insulin responses as well as the rate ofin vitro hydrolysis with a-amylase were strongly correlated to the degree ofstarch gelatinization (r = 0.88, r = 0.90, and r = 0.96, respectively). Plasma glucose and insulin responses were also positively correlated to the rate ofhydrolysis with a-amylase in vitro = 0.98 and r = 0.76, respectively). (r These results suggest that the degree of starch gelatinization is an important determinant both for thestarch rate of hydrolysis in vitro and for the metabolic response in vivo. Am J C/in Nuir 1988;47:
1010-6. KEY plasma WORDS insulin

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Starch

gelatinization,

starch

digestion

rate,

a-amybolysis,

plasma glucose,

tant factor in the rate ofstarch hydrolysis and metabolic response. in many common plant foods the starch is The variable glycemic response after ingestion of However, only partly gelatinized, because ofthe limited water constarchy foods has been the subject of much interest retent during processing. The starch granules are only cently, especially in relation to diabetes. So however, far, slightly swollen and the internal structure is partly intact. not much attention has been paid to the importance of Examples of such foods are breakfast flakes of cereal the degree ofstarch gelatinization (DG). grains (3) and several baked products (8- 1 1). The starch In plant cells starch is present as granules. Starch pobygranules in legumes may swell incompletely during promers (amybose and amybopectin) are tightly packed in cessing (12, 13). In view of the increased attention to granules with a high degree of molecular order and are starchy foods and the nutritional advantages carbohyof associated by hydrogen bonding. Raw granules contain drates that are slowly digested and absorbed (14-19), the highly crystalline regions and are birefringent in polarnutritional properties of incompletely gelatinized starch ized bight. The granules are insoluble in cold water. When exposed to heat in the presence ofwater, the starch are of interest. This paper describes the close relationship between the granules undergo an irreversible swelling and destruction DG of pure wheat starch and the enzymatic susceptibilof the internal crystalline structure and birefringence is as well as the glucose and insulin responses lost. This transformation is termed gelatinization. With ity in vitro to starch in rats. The results are discussed in terms of a excessive treatment the granules may even rupture and disintegrate and a fraction of the starch is then sobubilized. I From the departments ofFood Chemistry and Food Technology, Raw starch is only slowly digested by enzymes in vitro University of Lund, Lund, and the Department ofCell Biology, Urnwhereas cooking increases the susceptibility consideraversity ofLink#{246}ping, Link#{246}ping,Sweden. bly because ofthe rupture and disintegration ofthe com2 Address reprint requests to J#{246}rgen Holm, Department of Food pact crystalline granular structure (1-4). Furthermore, Chemistry, Chemical Center, University ofLund, P 0 Box 124, S-22l the glucose and insulin responses in vivo are significantly 00 Lund, Sweden. greater after ingestion of cooked compared with raw ReceivedFebruary2, 1987. Accepted for publication June 16, 1987. starches(3-7). Consequently, DG is an extremely impor1010
Am J C/in Nuir l988;47: 1010-6. Printed in USA. 1988 American Society Clinical for Nutrition

Introduction

STARCH
TABLE Physical 1 and

GELATINIZATION

AND

DIGESTION

RATE

bOll

morphological

characteristics

ofwheat-starch Degree Calorimetric method (DSC)[nJ % [3] Ot [3] l4.2 [4] 36.9t [5] 7 1 . 1 f [3] 85.0 [21 96.Of [2] l00 [2]

samples

inused the

experiments

of gelatinization

Treatment

temperaturet C
-

Heat

ofgelatinization J/g(cal/g)

En)

Enzymatic method [n] % 0. 1 0.4t 14. 1 47.3 91 .9 97.2 97.8f

-

Swelling power [nJ

Solubility [n] %

Microscopical appearance

10.52 47 50 53 56 59 65 9.02 6.64 3.04 1.59 0.42 0 [2]

0.29

(2.5 1 0.07) 0.04)[4) 0.06) [5] 0.01 ) [3] 0.0 1) [21

[4] [4] (6]

2.0 [4] 2.4 [4] 2.8 [3)

0.2 [4J 0.4 0. 1 [4J 0.6 0. 1 [3] 0.8 0. 1 [3] 1.4 0. 1 [3] 1.6 [3J 2.8 [3)

Intact

birefringent

0. 17 (2. 15 (1 .59 0.03 (0.73 0.02 (0.38 (0. 10) [2]

0.25

1 .3f 1.9f

0.4f
0.7f [2]

[4J
[2]

3.9 [31 5.4 0. 1 [3]

6. 1 0.2 [3] 6.9 0.2 [3]

granules Increased swelling and decreased number of granules showing birefringence Extensively swollen nonbirefringent granules Granules disintegrated

100 MeanSEM.

0 [2]

l00

[21

35.0 4.0(3J

Downloaded from www.ajcn.org by guest on August 28, 2012

t Partly gelatinized. Completely gelatinized.

physical starch. Materials Preparation and morphological characterization of sponding the
cording

t Raw.

to 20 mg of starch
to Chiang and Johnsson

was withdrawn
(21). DO was

and

analyzed sus-

acas the

expressed

percentage
ceptible

ofthe
and

total starch

content

that was immediately

to glucoarnylase. solubi/itypauerns

and

methods Swelling samples

The heat-treated suspensions were carefully transferred to Raw commercial wheat starch was obtained from KEBO weighed centrifuge bottles and diluted with distilled water to Lab AB (Stockholm, Sweden). The starch content was 98.2 g/ 30 g/L starch concentration. A suspension of raw starch was 100 g (polymer weight, dry basis), determined as glucose after kept for 20 mm at room temperature under gentle agitation. incubating the sample with a thermostable a-amylase at 95 #{176}C samples were centrifuged The for 15 mm 700 X g. The superat and amyloglucosidase at 60 #{176}C To obtain (20). starch with natant was decanted and evaporated to drynessin the oven and different DOs, 100 g/L suspensions ofwheat starch in distilled the amount ofsolute was measured. Granule swelling and soluwater were heated under gentle agitation at 47, 50, 53, 56, 59, bility were calculated according to Leach et al (22): or 65 C. After 20 mm the samples were cooled to room ternSwelling power perature. A boiled sample was prepared by boiling ag/L50 suspension for 20 mm. = weight ofgel/(total starch weight ofsolutes) (g/g) (2)
-

ofstarch

Degree

ofstarch

gelatinization

Solubility

(%)
=

The DO was measured by two different methods. Duplicate analyses were performed on each preparation. With the caborimetric method (differential scanning calorimetry [DSC]) we Microscopy measured the heat ofgelatinization (H) ofthe starch samples, A few drops
ie, the energy that has to be supplied to obtain complete bOO g/L suspension a starch microscopy

100

X weight

ofsolutes/total

starch

(gig)

(3)

acterize gelatinization. Ten to 15 mg of was heated from 22 to 80 #{176}C scanning at a rate of 10 C/min in in vitro coated sample pans. H was calculated from the peak area of Experiments the thermogram. The instrument used was a Perkin-Elmer A porcine pancreatic a-amylase preparation (27 mg protein/ DSC-2 (Perkin-Ebmer Corp, Eden Prairie, MN). The DG was mL, 1200 units/mg; Sigma Chemical Co, St Louis, MO) was calculated by comparing iH for raw starch with that for heat- diluted 1:20 or 1:400 before incubation. The heat-treated samtreated starch: ples were diluted with distilled water to 50 gIL. Forty-five milbiliters of0.05 mob/L Na,K-phosphate buffer(0.025 mol/L each DG (%) = (1 x 100 (1) of KH2PO4 and Na2HPO4) containing 0.4 NaC1 (pH 6.9) g/L The enzymatic method is based on the principle that gelatin- and 1.25 mL ofdiluted a-amylase preparation were added to a ized starch is easily digested by glucoamybase to form glucose. lO-mL subsample corresponding to 500 mg starch. Samples A small amount of the heat-treated starch suspensions correwere taken before and after 5-60 mm incubation (under gentle
-

10/L suspension g were examined by light light microscopy to further charthe morphology ofthe granules.
and bypolarized

ofa

1012 agitation at 37 #{176}C) analyzed and with dinitrosalicylic

HOLM
acid

ET AL
for of
70 80

content
using
starch

ofreducing
maltose.
degraded

sugar

(23).

A standard

curve

was prepared
proportion equivalents) equivalents

was calculated times 0.95 divided Experiments The

vivo in male

The extent of hydrolysis (the to maltose, or percent maltose as 100 times milligrams of maltose

by milligrams

ofstarch

in sample.
60

in vivo of starch
(Sprague-Dawley,

availability
rats

for digestion
120-150

and
g) was

absorption
studied by

in I)
Cl) >

50

analyzing

concentrations

of glucose

and

insulin

in plasma

at 0
40

various time intervals after gastricintubation. The starch samples were diluted with a NaCl solution to 40 gIL. The final con- I centration of NaCI was 9 gIL. After being starved for 24 h unanesthetized rats were intubated with a starch suspension (2.5 mL) corresponding to100 mg starch. Nine rats were used for
each sample tested. Blood was obtained from the rats by multipIe serial sampling from the retroorbital venous plexus. After 135 mm the rats were killed by a blow on the head and a piece of the liver (-0.5 g) was taken from the ventral part of the left lateral lobe for analysis ofglycogen content. Estimation of

30

20

Downloaded from www.ajcn.org by guest on August 28, 2012

plasma glucose concentration was done with a glucose-oxidase method (24). Plasma insulin levels (IRI) were analyzed with a radioirnmunoassay (25). Liver glycogen concentration was
determined St atistical by the evaluation method ofRerup and Lundquist (26).

10

0-

15

30

60

The

results

are given

as mean SEM.

The

significance

of

Incubation

time

(mm)

differences
Results

was tested

with Students t test.

The physical and morphological characterization the starch samples used in the in vitro and in vivo experiments are presented in Table 1. With increasing DG there was an increased susceptibility to glucoamylase as The boiled sample well as a gradual boss ofbirefringence, indicating that the sis rate, were still very pronounced. was hydrolyzed to 75% (maltose equivalents) within 5 highly ordered structure within the granules was dea plateau at 80% after 30 mm, whereas stroyed to different extents. The soluble starch fraction mm and reached was only slightly increased for the partly gelatinized sam- the hydrolysis values at 5 mm for raw starch and starch with DGs of 14 and 37% were b2, 30, and 48%, respecples and for the completely gelatinized sample heated at 65 #{176}C above (just the gelatinization temperature range). tively. course of the plasma glucose and insulin reIn contrast, after boiling, a large fraction of the starch The in rats after intubation with starch with different was soluble, indicating disintegration of the granular sponses DGs are shown inFigures 3 and 4 and the area under structure. are presented in Table 2. The plasma glucose The susceptibility of starch with different DGs to hy-the curves after intubation with boiled starch (DG drobysis by porcine pancreatic a-amylase (added at response a = 100%) was much greater than after raw starch (DG concentration of200 units/g starch where 1 unit liberates to the partly gelatinized products 1 mg maltose from soluble starch in 3 mm at pH 6.9 and = 0%). The responses The response to the partly gelatinized at 20 #{176}C) is shown in Figure 1. Raw starch was hydro- were intermediate. with a DG of 37% was greater than to that with a lyzed most slowly and the susceptibility to a-amylase in- starch DG of 14%. With raw starch the glucose peak dewas creased with DO. The two completely gelatinized sambayed (peak value at 60 mm), whereas the peak values ples(DG = 100%)and starch with a DG of96% displayed and completely gelatinized starch samples the highest availability; no significant difference was oh- for the partly were obtained after only 1 5 mm. The insulin responses served between these three samples. were closely associated with the glucose responses. The When a large excess ofenzyme was used by increasing glucose (0-60 mm) and plasma insulin (0the enzyme concentration 20-fold to 4050 units/g starch, rise in plasma 30 reflect the early rates of digestion and absorpthe initial hydrolysis rate increased as did the extent of mm) where the largest differences are found. The liver hydrolysis (Fig 2). However, the differences in availabil- tion, glycogen contents 1 35 mm after intubation were 4.1 ity between the samples, especially in the initial hydroly-

FIG 1. Hydrolysis ofstarch with different DOs by porcine pancreatic a-amylase added at a concentration of 200 units/g starch. Percentage hydrolysis is expressed in terms of maltose equivalents. DG (%): 0 (raw); A, 14.2; 36.9; 0, 71.1; t, 85.0. The bars represent the interval obtained with 96.0, lOO(heated at 65 C,just above gelatinization temof perature range), and 100%(boiled). The SEM did not exceed 3.4 for any experimental point. n = 2-3.

#{149},

#{149},

STARCH
90

GELATINIZATION

AND

DIGESTION

RATE

1013

80

70

60

I) I)

starch was hydrolyzed almost completely within 5 mm, barge differences in the rate of hydrolysis remained (Fig 2). Starch hydrolysis with a-amylase results in the formation ofglucose, maltose, maltotriose, andlimit a dextrins (branched obigosaccharides with four glucose monomers or more). Therefore, 75-80% hydrolysis expressed as maltose equivalents corresponds to an almost complete hydrolysis. Once the highly organized structures within the swollen granules were completely destroyed, as in the sample treatedjust above the gelatinization temperature range, the availability ofstarch was not increased further
by swelling, rupture, and structure (boiled sample) disintegration (Fig 1). The of the increased granular suscepti-

50

>

0
>

40

30

bility to a-amylase was not rebated to increased sobubility because the soluble starch fraction did not exceed 3% in any sample, except for boiled starch. The plasma glucose and insulin concentrations in rats were raised to different extents depending on the DG. An
already bow DG (14%), which caused only slight swelling

Downloaded from www.ajcn.org by guest on August 28, 2012

20

10

and internal disorganization of the granules (Table 1), raised the glycemic response far above that ofraw starch. The decline in plasma glucose concentration was slower for partly gelatinized starch than for completely gelatin-

0
I

___0

15

30

60 _i

smgnifmcant . . .
A-I A1-0 A U 0
p

dmfferences
aQ

(mm)
0.001

Incubatmon

tmme

(1Tn)

<

0.00 0.00

1 1 p

< < <

p 0.001
1 1

p <

p 0.00
0.00 0.001

p p

<

FIG 2. Hydrolysis ofstarch with different DGs using a large excess of porcine pancreatic a-amylase (4050 units/g starch). DG (%): 0 (raw); A, 14.2; 36.9; 0, 100 (boiled). The SEM did not exceed 3.4 for any experimental point. n = 2-3.

#{149},
10
-I

<

0.00 1
0001

o.oo
0.05

p<0.01

p<

p<

p<

p<o.o5

0.7 (raw), 6.9 0.8 (DG= 14%), 6.1 1.1 (DG = 37%), and 6.3 0.5 (boiled) mg/g wet wt ofliver. The glycogen content was significantly bower with raw starch than with boiled starch or starch with DG of (p14%
< 0.05).

9,

E E
0)
Cl)

The

liver

glycogen

content

in control

rats(n 4) =

0 intubated with 0.9% NaCb solution was 0.1. 0.3 C) The plasma glucose and insulin responses were posi- a) tiveby correlated with the rate of hydrolysis with aE amylase in vitro, and both the plasma glucose and insu- (0 CO lin responses as well as the rate ofhydrolysis with a-amy- 0 lase were positively correlated with the DG (Table 3).

Discussion
3, During gelatinization interand intramolecular hydrogen bonds are broken. This results in a loosening up ___________________________________ 1#{149} I I I I of the compact granular structure and allows different 0 15 30 60 120 degrees of swelling and absorption of water, fully hydrated starch molecules leach from the granule. ConseTime after intubation (mm) quently, the availability of the starch granules digesto FIG 3. Plasma glucose responses at various intervals in rats given tive enzymes increases to different levels with increasing h with different DOs (100 mg as a 40 g/L suspension in 9 g/L DG. Even when we used an enzyme concentration in our DG (%): NaCI). 0 (raw); A, 14.2; 36.9; 0, 100 (boiled). Nine rats in vitro system that was so high that the boiled wheat were used for each sample tested. (Mean SEM.)

#{149},

#{149},

1014

HOLM

ET AL
TABLE 3 Correlation

significant
i.
#{149}-A #{149}-o p<0.05 p<0.01

differences
122

coefficients between
and plasma

DO,

digestion

(mm)

and plasma glucose

insulin responses

rate ofstarch in rats

in vitro,

r
p<0.01

A-D 0.3 1-0

p<0.05

-a 0
0.2

DO with digestion rate(n = 7)t DO with plasma glucose(n = 4) DO with plasma insulin (n = 4) Digestion rate with plasma glucose (n Digestion rate with plasma insulin (n

= =

4) 4)

0.96 0.88 0.90 0.98 0.76 ofthe heat of after 60 m m of 200 units/g Plasma insuon which the

E
C

a:

* DO was calculated from calorimetric measurement starch gelatinization. Digestion rate = degree ofhydrolysis incubation with a-amylase (added at a concentration starch). Plasma glucose = area under the curve, 0-60 mm. lin = area under the curve, 0-30 mm.

t n

number were

of starch
based.

samples

with different

DGs

calculations
0

provide glucose with prolonged absorption in the treatment of type 1 glycogenosis (27). The effect of raw 120 0 15 30 60 starchy foods on the glycemic response may be more favorable than supplementation with gel-forming types of Time after intubation (mm) dietary fiber. The use of raw plant food in the nutrition FIG 4. Plasma insulin (IRI) responses at various intervals in rats ofdiabetic patients was discussed (28). given starch with different DOs (100 mg as a 40 g/L suspension in 9 Despite a low enzymatic availability, raw wheat starch g/L NaG). DG (%): 0 (raw); A, 14.2; I, 36.9; 0, 100 (boiled). Nine is almost completely digested and absorbed in the rat rats were used for each sample tested. (Mean SEM.) small intestine (29). Raw potato starch, on the other hand, is poorly digested (30). In some processed food products the starch is only ized starch, which confirms that digestion and absorppartly gelatinized with a slightly swollen granular struction are prolonged when swelling is limited. Raw starch granular organization partly intact. was absorbed very slowly and elicited a flattened plasma ture and the internal For example, some baked products and breakfast cereals glucose curve with a delayed and less pronounced peak contain incompletely gelatinized starch granules, which and a very slow decline. is attributed to the limited water available (10-60 g/lOO The effect ofcompbete gelatinization on metabolic reg) when these products are produced. Cereal flakes are sponse has been studied in humans, both in diabetic and produced by steaming whole kernels and then flaking nondiabetic subjects(5-7)and in animals(3, 4). Potatoes them between rollers. Starch in breakfast cereals or preand corn starch give much lower postprandial glucose cooked convenience foods produced under more severe and insulin responses in raw form than after cooking (5conditions (elevated temperatures, pressures, and shear 7). In fact, raw corn starch has been used clinically to forces, eg, extrusion cooking or popping) is normally completely gelatinized despite the bow water content. A recent study (3) found that the plasma glucose and insuTABLE 2 bin responses in rats were lower after ingestion of wheat Area under the curve after gastric intubation ofrats with starch with flakes with a DG of45%, as measured by DSC, than after different DG5 boiled completely gelatinized whole-grain wheat flour. Area under the curve The in vivo responses were closely related to the in vitro digestibility with pepsin and a-amylase. Plasma insulin Plasma glucose Plasma glucoset According to Wootton and Chaudhry (1 1) DGs in (0-120 mm) (0-30 mm) DO (0-60 mm) short bread, hard sweet cookies, soda crackers, crispnmo!/L X mm % mmol/L X mm mmol/L X mm bread, wafer, fruit cake, and bread crumbs were 2, 3, 1 -0.310.66 28227 0 l26l333, 40, 50, and 60%, respectively. The corresponding in 0.350.53 14 r2lllOIt 138119 t vitro digestibility values with porcine pancreatic af-0.250.55 146522 t 37 11124815 It amybase were 18, 25, 33, 43, 56, 66, and 69%, respec1 2.120.68 100 1t-293l1Jt t151225 t tively whereas pregelatinized wheat starch was digested Mean 5EM; n = 9. For each bracket, values without footnotes were sig. to 90%. The variations in gelatinization were explained nificantly different from values with footnotes. in terms ofprebaking water content, baking time at high t < 0.001. moisture bevel favoring high DG, and the presence of t p <0.01. added ingredients that restrict gelatinization. Lineback p<0.O5.
I I I I

Downloaded from www.ajcn.org by guest on August 28, 2012

#{149},

STARCH

GELATINIZATION

AND

DIGESTION

RATE

1015

and Wongsrikasem (10) found DOs between 4 and 97% closely correlated (r = 0.99) with those obtained with in several commercial U.S. baked products. Hagander et DSC (Table 1). The characterization ofthe starch, especially regarding al (3 1) observed that extruded crispbread elicited larger the degree ofgebatinization, would be desirable in studies glucose and insulin responses than a conventionally with the rate of digestion and absorption of baked bread in diabetic patients, possibly because theconcerned starchy foods. Light microscopy in combination with poconventionally baked bread had a bower DG. barized light microscopy as webb as enzymatic methods Snow and ODea (1) obtained a more rapid hydrolysis used in this study are rapid methods that are of starch in vitro in commercial breads compared with like that for this purpose. homemade breads and suggested that this might have well suited In conclusion, the rate ofstarch hydrolysis in vitro and been due to a greater exposure to heat in the commercial and insulin responses in vivo are increased to breadmaking process. L#{252}der al (32) reported et signifi- the glucose extents, depending on DO. Hence, starchy cantby lower blood glucose bevels in healthy subjects after different with low DOs may be beneficial nutritionally beingestion of 85% rye-15% wheat bread with a baking foods cause they favor a reduced rate ofstarch uptake. 13 time of35 mm compared with bread with a baking time of 45, 55, or 65 mm. These differences in glycemic response were probably caused by differences in DO that References were caused by the variations in baking time. The extent ofstarch gelatinization also varies within a the rate ofhydrolysis of starch given bakery product because ofa moisture gradient (8). 1. Snow P, ODea K. Factors affecting in food. Am J Gin Nutr 198l;34:272l-7. Thus, starch gelatinization in white pan bread ranged 2. El Faki HA, Venkataraman LV, Desikachar HSR. Effect of profrom 33% in the crust to 70% in the center ofthe crumb cessing on the in vitro digestibility of proteins and carbohydrates (8). The crust contained many unswollen birefnngent in some Indian legumes. Qual Plant 1984; 34:127-33. starch granules whereas starch granules in the crumb 3. Holm J, Bj#{246}rckAsp N-G, Sj#{246}berg Lundquist I, L-B, I. Starch availcenter were deformed and only slightly birefringent. The ability in vitro and in vivo after flaking, steam-cooking and popping ofwheat. J Cereal 5th 1985;3: 193-206. differences in DG between crumb and crust is the proba4. Lee PC, Brooks SP, Kim 0, Heitlinger LA, Lebenthal E. Digestibilble reason for differences in postprandial hyperglycemia ity of native and modified starches: in vitro studies with human after ingestion ofbread with different crust-to-crumb raand rabbit pancreatic amylases and in vivo studies in rabbits. J tios (33). Bread rolls with high crust-to-crumb ratios Nutr 1985; 1 15:93-103. raised the blood glucose level more slowly with a later 5. Collings P, Williams C, MacDonald I. Effects ofcooking on serum peak and a slower decline than loaves with bow crust-toglucose and insulin responses to starch. Br ed J 198 l;282: M 1032. crumb ratios. 6. Vaaler 5, Hanssen KF, Aagenaes 0. The effect ofcooking upon the Dreher et al (34) obtained a higher in vitro digestibility blood glucose response to ingested cai-rots and potatoes. Diabetes and more extensive gelatinization of starch in potato Care l984;7:221-3. J, Sidbury JB. A study ofabsorption of differchips compared with starch in baked potato. The in vitro 7. Chen YT, Muenzer ent starches with equivalent as cooked food. Gin Res l983;3l: digestibility of potato chips, baked potato, and raw po6 15A(abstr). tato were 66.3, 53.6, and 228%, respectively. 8. Varriano-Marston E, Huang VKEG, Ponte J. Comparison of Differences in the extent of gelatinization after promethods to determine starch gelatinization in bakery foods. Cereal cessing legumes in different ways was suggested as a plauChem l980;57:242-8. sible explanation for differences in in vitro starch digest- 9. Wootton M, Weeden D, Munk N. A rapid method for the estimaibility (2). It has been suggested that the low metabolic tion ofstarch gelatinization in processed foods. Food Technol Aust response and in vitro starch digestibility of legumes is 197l;23:6l2-5. 10. DR. Wongsrikasem E. Gelatinization ofstarch in baked caused by the entrapment of starch in the cells (12, 13, Lineback products. J Food Sci1980;45:71-4. 35). The cell walls may limit the hydrolysis rate because M, Chaudhry MA. Gelatinization and in vitro digestibilof steric hindrance to enzymatic attack (13). Further- 1 1. Wootton ity ofstarch inbaked products. J Food 5th980;45:l783-4. l more, the cell walls may also inhibit starch gelatinization 12. Thorne Mi, Thompson LU, Jenkins DJA. Factors affecting starch by physically limiting the degree ofswelling ofthe starch digestibility and the glycemic response with special reference to legranules and by limiting the water transport necessary gumes. Am J Clin Nutr l983;38:48l-8. for complete gelatinization (12, 13). Grinding raw be- W#{252}rsch Del Vedovo 13. P. S, Koellreutter B. Cell structure and starch gumes, thus disrupting the cell walls, followed by cooknatureas keydeterminantsofthedigestion rate ofstarch in legume. Am J Gin Nutr 1986;43:25-9. ing resulted in an increased swelling ofthe granules, and M, Kolterman 0G. Comparison of seit was proposed that this could increase the rate of starch 14. Crapo PA, Insel J, Sperling rum glucose, insulin, and glucagon responses to different types of digestion as well as the glycemic response (12, 13). complex carbohydrate in noninsulin-dependent diabetic patients. DSC is a calorimetric method based on the physical AmJClinNutr l98l;34:l84-90. course of gelatinization but it requires equipment not15. Jenkins DJA, Wolever TMS, Taylor RH, et al. Glycemic index of usually available in most nutritional laboratories. Howfoods: a physiological basis for carbohydrate exchange. Am J Clin ever, the values for DU obtained with the simple and Nutr l98l;34:362-6. rapid enzymatic method using glucoamylase were16. Jenkins DJA, WoleverTMS, Kalmusky J. Lowglycemic index car-

Downloaded from www.ajcn.org by guest on August 28, 2012

1016
bohydrate foods in the Nutr l985;42:604-l7.
17. Brudevold F, Goulet management D, Tehrani ofhyperlipidemia. Am

HOLM
J Can

ET AL
26. RerupC, Lundquistl. Non-specific reaction ofcurrentglucose oxidase preparations with glycogen and application its for glycogen determinations in tissue. Acta Pharmacol Toxicol l967;25:4l-53. Wolfsdorf JI, Laffel LB, Crigler JF Jr. Uncooked cornstarch: requirements for biochemical control Type in I glycogenosis. Gin Res 1984; 32:764A(abstr). DougiassJM. Raw diet and insulin requirements. Ann Intern Med 1975;82:6 1-2. Bj#{246}rck Nyman I, M, Pedersen B, Siljestrom M, Asp N-G, Eggum BO. On the digestibility ofstarch in wheat bread-studies in vitro andinvivo.JCerealSci 1986;4:l-ll. Holm J, Bj#{246}rck Ostrowska I, S, et al. Digestibility of amylose-lipid
complexes in-vitro and in-vivo. Starch/St#{228}rke l983;35:294-7.

A, Attarzadeh F, van J. Houte and maltose clearence from wheat 27. 19:136-44. ofglucose tolerance and plasma insulin to 28. the incidence of coronary heart disease: results from two opulap tion studies in Finland. Diabetes Care l979;2:l31-4l. 19. Heaton KW. Fibre, satiety and insulin-a new approach to over- 29. nutrition and obesity. In: Heaton KW, ed. Dietary fibre: current developments ofimportance to health. London: John Libbey & Co

Intraoral demineralization starch.CariesRes 1985; 18. Pyorala K. Relationship

Ltd. 1978:141-8.

30.

20. 21.

22.

23.

24. 25.

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