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Starch digestibility in food matrix: a review.

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 Author's personal copy
Trends in Food Science & Technology 21 (2010) 168e180
Starch digestibility in
food matrix: a review
Jaspreet Singh*, Anne Dartois
and Lovedeep Kaur
Riddet Institute, Massey University,
Palmerston North, New Zealand
(Tel.: D64 6 3505062; e-mail: j.x.singh@massey.ac.nz)
In this review, recent reports on in vitro starch hydrolysis kinet-
ics are reviewed with regard to the structural characteristics of
starches. Factors such as starch granule morphology, amylose
to amylopectin ratio, molecular structure, degree of branching
in terms of steric hindrance and consequently mass transfer re-
sistance and their effects on the digestibility and absorption of
digested carbohydrates have been examined. The physical
state of the starch ingested has a major impact on the digest-
ibility therefore effects of processing techniques (thermal pro-
cessing, extrusion cooking, autoclaving etc.) and starch
modification have been discussed. The other constituents of
the food matrix, such as proteins, lipids and polysaccharides,
play a significant role during processing which affects the
physico-chemical characteristics of digesta and the final di-
gestibility of starch. Some molecules naturally occurring in
food sources may confer an inhibitory effect during starch
hydrolysis.
Introduction
Starch is the commonest storage carbohydrate in plants
and also the largest source of carbohydrates in human
food. Starch consist of two types of molecules: amylose (lin-
ear polymer of a-D-glucose units linked by a-1,4 glycosidic
linkages) and amylopectin (branched polymer of a-D-glu-
cose units linked by a-1,4 and a-1,6 glycosidic linkages).
Starch and starchy food products can be classified according
to their digestibility, which is generally characterized by the
rate and the duration of the glycemic response. Predicting
and controlling the glucose absorption due to ingestion of
starchy food is of great interest in the context of worldwide
health concerns. Most starches contain a portion that digests
* Corresponding author.
0924-2244/$ - see front matter Ó 2009 Elsevier Ltd. All rights reserved.
doi:10.1016/j.tifs.2009.12.001
Review
rapidly (rapidly digesting starch), a portion that digests
slowly (slowly digesting starch) and a portion that is resis-
tant to digestion (resistant starch) (Englyst, Englyst, Hudson,
Cole, & Cummings, 1999). Resistant starch has been defined
as the portion of starch that is not hydrolysed by the enzymes
in the small intestine and passes to the large intestine. One of
the most widely used methods to classify the starches based
on kinetics of in vitro digestion was suggested by Englyst,
Kingman, and Cummings (1992). The method is based on
the in vitro digestion of starch by simulating stomach and in-
testinal conditions and measuring glucose release at differ-
ent times. Based on the method, different starch fractions
are defined as:
e Rapidly digestible starch (RDS): amount of glucose
release after 20 min.
e Slowly digestible starch (SDS): amount of glucose
released between 20 and 120 min of in vitro digestion.
e Resistant starch (RS): total starch minus amount of glu-
cose released within 120 min of in vitro digestion
which can be explained by the following equation:
RS ¼ TS  ðRDS þ SDSÞ ð1Þ
where TS is total starch.
Accurate determination of bioavailable carbohydrate in
a given product enables the manufacturer to communicate
the glycemic response per serving of a food, especially in
the case of diabetes therapeutic food, management of dia-
betes and disorders of carbohydrate metabolism. The con-
cept of glycemic index has been introduced to classify
foods on the basis of their postprandial blood glucose re-
sponse. The glycemic index is defined as the postprandial
incremental glycemic area after a test meal, expressed as
the percentage of the corresponding area after an equi-car-
bohydrate portion of a reference food such as glucose or
white bread (Goni, Garcia-Alonso, & Saura-Calixto,
1997; Jenkins et al., 1987). Several workers have indicated
a good relationship between the rate of in vitro digestion
and the glycemic response to food. Such studies may be
used to identify food of potential use in the diet of individ-
uals with diabetes.
Physico-chemical and structural characteristics of starch
vary among different botanical sources (Singh, Kaur, &
McCarthy, 2007; Singh, Kaur, & McCarthy, 2009). Several
studies have indicated relationships between different
 Author's personal copy
J. Singh et al. / Trends in Food Science & Technology 21 (2010) 168e180
starch characteristics and in vitro digestibility. Chemically
modified starches are commonly used in food applications
as they offer improved functional properties than native
starches. Several reports suggest that chemically modified
starches exhibit greater resistance towards a-amylase diges-
tion (Han & BeMiller, 2007). The textural and rheological
characteristics of food; the presence of other food compo-
nents, such as proteins, lipids and non-starch polysaccha-
rides; and the changes and interactions occurring in them
during food processing affect the enzymatic digestibility
of starch to a considerable extent. Our review presents a de-
tailed survey on the factors affecting the starch digestibility
in food. It includes all the scales, from starch structure,
food processing to the digestion, with a particular focus
on the effects of other components present in the food ma-
trix. The recent information about in vitro enzymatic di-
gestibility and the factors affecting it has been analyzed
and is presented in this review.
Kinetics of starch digestibility
The digestible starch is mainly hydrolyzed by the en-
zymes into glucose through several steps. Salivary a-amy-
lase acts quite efficiently on starch but is rapidly degraded
in the acidic environment of the stomach and hence plays
a very minor role in the process of starch digestion. The
majority of the starch hydrolysis is carried out by the pan-
creatic amylase, which is released in the small intestine via
pancreatic duct. a-Amylase (1,4 a-D-glucanohydrolase, EC
3.2.1.1) catalyzes the hydrolysis (endo attack) of a-(1e4)
glycosidic bonds in amylose and amylopectin of starch
(Lehmann & Robin, 2007). Both the linear and branched
(amylose and amylopectin) polymers of starch are hydro-
lysed by virtue of binding of their five glucose residues ad-
jacent to the terminal reducing glucose unit to specific
catalytic subsites of the a-amylase, followed by cleavage
between the second and third a-1,4-linked glucosyl residue
(Gray, 1992). The final hydrolysis products from amylose
digestion are mainly maltose, maltotriose and maltote-
traose. However, a-amylase from some microbial sources
may produce maltohexose and maltoheptose along with
maltotriose (Yook & Robyt, 2002). a-Amylases have no
specificity for a-1,6 branch linkage in amylopectin, there-
fore their capacity to break a-1,4 links adjacent to the
branching point is decreased mainly due to steric hin-
drance. The results obtained from the analysis of intestinal
content of humans suggest that hydrolysis products from
amylopectin mainly consist of dextrins or branched oligo-
saccharides. The products obtained from a-amylase starch
hydrolysis have been observed to possess a-anomeric con-
figuration of the substrate (Kuriki & Imanaka, 1999). The
resulting oligosaccharides (maltose, maltotriose and a-dex-
trins) are further hydrolyzed efficiently by the action of
brush border enzymes of the intestine.
Several in vitro methods have been proposed in the past
by various researchers to evaluate the rate of starch digest-
ibility through hydrolysis. Many of these methods vary in
169
regard to sample preparation especially cooking procedures,
use of amylases and proteases and the confirmation of find-
ings through an in vivo method. An account of some recent
studies on starch digestibility is given in Table 1. The pro-
cess of in vitro starch hydrolysis is highly influenced by
starch characteristics and composition of starch based
foods. There is a wide variation in the rate of starch hydro-
lysis among different food products, from the lowest for len-
tils, and to the highest for boiled potatoes. An improved in
vitro method to study starch hydrolysis at different time in-
tervals was suggested by Goni et al. (1997). A first order
equation for the hydrolytic process of starch hydrolysis in
foods has been proposed:


C ¼ CN 1  ekt ð2Þ
where C is the concentration of starch hydrolyzed at time t,
CN is the equilibrium concentration, k is the kinetic con-
stant and t is the chosen time. By using this equation, it
has been observed that legumes are hydrolysed to a much
lesser extent than cereals. The term hydrolysis index was
also suggested by Goni et al. (1997) based on the relation-
ship between area under the hydrolysis curve (AUC) for
a given sample and the AUC for a reference food such as
white bread. The authors suggested that the hydrolysis in-
dex can be a good predictor of glycemic response. The
AUC is generally calculated using the following equation:

 

AUC ¼ CN tf  to  ðCN =kÞ 1  exp  k tf  to
ð3Þ
where CN corresponds to the equilibrium percentage of
starch hydrolyzed after 180 min, tf is the final time
(180 min), to is the initial time (0 min) and k is the kinetic
constant. Kinetics at low starch concentrations can also be
described through a simple MichaeliseMenten equation
whereas a modified first-order MichaeliseMenten model
is required at high concentrations (Komolprasent & Ofoli,
1991). MichaeliseMenten kinetics describes the velocity
of liberation of reducing sugars as a function of initial
starch concentration, although the concentration of a-1,6
branching also affects it to some extent. The maximal ve-
locity (Vmax) of the liberation of reducing sugars from
starch decreases whereas the value of the Michaelise
Menten constant (Km) increases with decreasing molecular
weight of starches (Heitmann, Wenzig, & Mersmann,
1997). Maltose produced during the hydrolysis may inhibit
a-amylase starch hydrolysis noncompetitively, although an
uncompetitive inhibition by glucose and maltose has been
reported recently for a-amylase hydrolysis of rice starch
(Apar and Özbek, 2007). The structural differences among
the starches, such as molecular weight and degree of
branching, may influence the rheological characteristics
of their solutions which also affect the velocity of starch
hydrolysis considerably.
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170 J. Singh et al. / Trends in Food Science & Technology 21 (2010) 168e180

Table 1. Digestibility of different starches and starch based foods.

Source Starch digestibility Reference and method

a
Cooked potatoes >90 Mishra et al. (2008); in vitro simulated gastric and small intestinal
digestion using pancreatin and amylogucosidase
Sorghum meal 60e85b and 40e47c Wong et al. (2009); in vitro starch digestion with pepsin and without
pepsin pre-treatment using pepsin (porcine stomach mucosa) and a-
amylase (bacterial; porcine pancreas; human saliva)
Cooked rice noodle dough 43d and 33d Koh et al. (2009); in vitro digestion using a-amylase from
Aspergillus oryzae
Waxy maize starch 100e Han and BeMiller (2007); in vitro digestion by the method of Englyst et al.
(1999) using pepsin, pancreatin and amyloglucosidase
Chemically modified 76e87e
waxy maize starch
Normal maize starch 99e
Chemically modified 71e96e
normal maize starch
Potato starch 96e
Chemically modified potato starch 67e76e
Wheat flour 72f Englyst et al. (1999); in vitro digestion using pepsin, pancreatin
and amyloglucosidase
Corn flakes 81f
Cooked rice 70e80g Frei et al. (2003); in vitro digestion by the method of Goni et al. (1997)
using pepsin, a-amylase and amyloglucosidase from Aspergillus niger
Extruded amaranth seeds 93h Capriles et al. (2008); in vitro digestion by the method of Goni et al.
(1997) using pepsin, a-amylase and amyloglucosidase
Amylose-lipid complexes 48e71i Crowe et al. (2000); in vitro digestion using a-amylase and
amyloglucosidase
Legume starches 80e90j Hoover and Zhou (2003); treatment
with porcine pancreatic a-amylase
Extruded beans 290e306k Alonso et al. (2000); in vitro digestion with pancreatic amylase
Autoclaved legumes 87e89l Rehman and Shah (2005); in vitro digestion with pancreatic a-amylase
a
Expressed as rapidly and slowly digestible starch (%).
b
Expressed as mg reducing sugar/h (with pepsin pre-treatment).
c
Expressed as mg reducing sugar/h (without pepsin pre-treatment).
d
Expressed as mg of maltose equivalent liberation per g of dough.
e
Expressed as digestibility (%).
f
Expressed as total glucose after 120 min incubation with enzymes.
g
Expressed as digestible starch (%).
h
Expressed as hydrolysis index.
i
Expressed as conversion to glucose (%).
j
Expressed as hydrolysis (%).
k
Expressed as starch digestibility (mg of maltose g-1).
l
Expressed as starch digestibility (%).

Sanromán, Murado, & Lema (1996) studied the influ-
ence of starch structure on the kinetics of glucoamylase hy-
drolysis using two types of starches differing in their
apparent viscosity. In one case, the authors found that the
starch hydrolysis kinetics corresponds to Michaelise
Menten behaviour:
ðV ¼ Vmax S=ðKm þ SÞÞ
while for the other case, the hydrolysis rate of starch in-
creased with the concentration up to a maximum and
then a decrease was observed once a determined concentra-
tion was surpassed. In the later case an inhibitory effect of
the substrate (S ) due to its higher apparent viscosity was
postulated in the model
ðV ¼ Vmax S=ðKm þ SþKi SÞÞ ð5Þ
where Ki is the inhibition constant. The mass transfer limi-
tation occurs due to the viscosity/rheological characteristics
of the starch, which may also influence the hydrolysis rate of
different starches. A higher degree of branching allows an
increase in the number of available points for the enzymatic
ð4Þ attack (Sanromán et al., 1996). However, the branching may
increase the steric hindrance and, consequently, the mass
transfer resistance. This explains the behaviour of starches
at higher concentrations when the positive effect of branch-
ing is generally overcome by the diffusional restrictions to-
wards the enzymatic action due to higher apparent viscosity.
Therefore a decrease in starch hydrolysis may be observed
because of mass transfer limitation. In addition, the
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J. Singh et al. / Trends in Food Science & Technology 21 (2010) 168e180 171

molecular weight distribution of the starches may also affect Table 2. Starch characteristics and their effect on starch
the enzymatic hydrolysis. Indeed a higher molecular weight digestibility.
can result in an increased hindrance towards active centres Starch characteristic Starch Reference
of the enzyme. The kinetics of the hydrolysis reaction de- digestibility
scribed through the MichaeliseMenten equation suggested Granule morphology
that a Km value is influenced by the degree of polymerization Native potato starch 0.86a Kaur et al. (2007)
and content of a-1,6 branching bonds of the starches (Heit- Large potato starch granules 0.83a
mann et al., 1997). All these studies confirm that the rate of Medium potato starch granules 1.12a
starch hydrolysis is controlled by mass transfer rate and the Small potato starch granules 1.35a
Amylose content
effects of the starch structure are dependent on the substrate Waxy maize starch (cooked) 96b Wolf et al. (1999)
concentration. Normal maize starch (cooked) 86b
High amylose maize starch 77b
Starch characteristics and starch digestibility (cooked)
Waxy rice (cooked) 80c Hu et al. (2004)
The morphological characteristics of starches from var-
Normal rice (cooked) 71c
ious botanical sources vary with the genotype. The varia- Waxy sorghum starch (raw) 90d Sang, Bean, Seib,
tion in the morphology such as size and shape of starch Pedersch, & Shi
granules is attributed to the biological origin (Singh, (2008)
McCarthy, & Singh, 2006, Singh et al., 2007). Firstly, Normal sorghum starch (raw) 81d
Langworthy and Deuel (1922) reported an apparent direct a
Expressed as hydrolysis rate (%) of starch granules.
b
negative relationship between large size granules and starch Expressed as digestible starch (%).
c
digestibility. Many studies were carried out afterwards to Expressed as digestible starch (%).
d
Expressed as rapidly and slowly digestible starch (%).
validate this relationship. Lindeboom et al. (2004) reported
that the small barley and wheat starch granules hydrolyze
faster than the large granules. Kaur, Singh, McCarthy, &
Singh (2007) observed that significant differences exist
among the enzymatic hydrolysis values for different native sources has been reported by Dreher, Dreher, & Berry
potato starches and their separated small, medium and large (1984). The authors suggested a higher digestibility for
fractions (Table 2). Among native starches, potato starch the cereal starches compared to tuber and legume starches.
with a higher percentage of small granules showed a higher This may be attributed to the presence of numerous pin-
hydrolysis rate than those containing large or medium size holes on the surface layer and pores that penetrates towards
granules. The separated large granule fraction showed the interior of the granules from cereal sources, such as
a lower hydrolysis rate than medium and small granule maize. The pores in the granule facilitate the entry of the
fractions. The lower susceptibility of the large granule amylases for digestion. This type of hydrolytic attack
starches to enzymatic hydrolysis has been suggested to be may be termed endocorrosion. The sites of enzymatic
due to their smaller granule specific surface area, which action on the starch granules have also been reportedly
may decrease the extent of enzyme binding and ultimately influenced by granule source. Starch from normal maize,
result in less hydrolysis than small granules (Tester et al., sorghum and millet form random round pits on the granule
2006). Capriles, Coelho, Guerra-Matias, & Areas (2008) surface followed by internal hydrolysis while waxy cereal
studied the digestibility of amaranth starch and their results starches show point hydrolysis (Dreher et al., 1984). Native
suggested that the higher in vitro digestibility of amaranth or uncooked potato starch granules due to their smooth sur-
starch is attributed to its small granule size distribution face and the absence of any pores or pits have been
ranging between 1 and 3 mm. observed to have very high resistance towards enzymatic
The surface of the starch granules from potatoes corn, hydrolysis. First studies on the enzymatic resistance proper-
rice and wheat has been reported to be smoother than ties of uncooked potato starches were reported by Lang-
corn, rice and wheat starch granules. Li, Vasanthan, Ross- worthy and Deuel (1920) using human subjects. After
nagel, & Hoover (2001) observed the presence of ‘pin that many studies reported the digestibility and fermenta-
holes’ and equatorial grooves or furrows in large-sized tion of uncooked potato starches (Tester et al., 2006).
corn starch granules. The presence of small nodules and Potato starches are generally eroded by exocorrosion (ero-
protuberances on potato starch granule surfaces, ranging sion of the entire granule surface or sections of it) during
up to 1 mm in size, has been reported in the literature. Singh enzymatic hydrolysis.
et al. (2006) have also reported the presence of some small The presence of some non-starchy substances such pro-
nodules or protuberances, and surface fragmentation, on teins over the granule surface may also limit the rate of en-
potato starch granules. The surface characteristics of the zymatic hydrolysis. Granule surface proteins and lipids can
starch granules have been observed to affect their enzy- reduce surface accessibility by blocking the adsorption sites
matic digestion. A comprehensive account of the enzymatic and therefore influences enzyme binding (Oates, 1997).
digestibility of native uncooked starches from different The individual granules, in the case of rice starch, develop
 Author's personal copy
172 J. Singh et al. / Trends in Food Science & Technology 21 (2010) 168e180
into compact spherical bundles or clusters, known as com-
pound granules, which fill most of the central space within
the endosperm cells. To initiate the hydrolytic attack, the
amylases must functionally bind the glucan chains via sev-
eral glucose units to their subsites, which are located at the
active centre and are capable of interacting with one glu-
cose residue in the substrate. The number of active sites
within an active centre vary from 4 to 9. The ‘A’, ‘B’,
and ‘C’ patterns are the different polymeric forms of the
starch that differ in the packing of the amylopectin double
helices. The molecular structure of the starch granules es-
pecially the arrangement in A-type or B-type crystallites in-
fluence their hydrolysis pattern. A-type polymorphic
residues are less resistant to amylase hydrolysis than
B-type. The shorter double helices and interior crystallites
present in A-type starches are more susceptible whereas
longer chains form longer and more stable helices and are
more resistant towards enzymatic hydrolysis (Lehmann &
Robin, 2007).
The amylose content of the starch granules varies with
the botanical source of the starch which may be attributed
to differences in the activities of enzymes involved in the
biosynthesis of linear and branched components. Amylose
content of the starches varies from 14 to 31% for normal
genotypes. However waxy genotypes, essentially without
amylose, and high amylose starches having amylose up to
70% have also been reported. Raw starches high in amylo-
pectin have been shown to digest more quickly than those
high in amylose. Legumes contain 30e40% amylose and
60e70% amylopectin in their starch granules, while most
other food starches contain 25e30% amylose and
70e75% amylopectin (Hoover & Zhou, 2003). Amylopec-
tin is a much larger molecule than amylose with average
molecular weight at 105 and 106, whereas the average mo-
lecular weight for amylose is 104. Therefore amylopectin
has a much larger surface area per molecule than amylose,
which makes it a preferable substrate for amylolytic attack.
Indeed, raw legume starch with high amylose has been
shown to be less digestible than corn starch in rats, and
the rate of hydrolysis of legumes starch in vitro is less
than that of corn starch (Thorne, Thompson, & Jenkins,
1983). In addition, legumes contain twice as much protein
as cereals and anti-nutrients, which may affect the starch
digestibility. Furthermore, the glucose chains of amylose
starch are more bound to each other by hydrogen bonds
making them less available for amylolytic attack than am-
ylopectin which has many branched chains of glucose.
For these reasons the nature of the starch in carbohydrate
foods is important to consider the expected glycemic re-
sponse. Higher amylose content lowers the starch digest-
ibility because a positive correlation exists between
amylose content and resistant starch formation.
The postprandial glycemic and insulinemic responses of
some commonly consumed maize products were studied
and a higher resistant starch content was estimated for the
products containing high amylose maize starch (Granfeldt,
Drews, & Björck, 1995). Similarly, Frei, Siddhuraju, &
Becker (2003) and Hu, Zhao, Duan, Linlin, & Wu (2004)
reported the in vitro starch digestibility of different rice cul-
tivars varying in amylose content. Their study suggested
that starch hydrolysis was more rapid and more complete
for waxy cultivars than for the high amylose rice cultivars
(Table 2).
Phosphorus is one of the non-carbohydrate constituents
present in the starches, which significantly affects the func-
tional properties of the starches (Singh, McCarthy, Singh,
& Moughan, 2008). Phosphorus is present as phosphate
monoesters and phospholipids in various starches. Phospho-
lipids present in starch have a tendency to form a complex
with amylose and long branched chains of amylopectin,
which results in limited swelling. Studies on the enzymatic
and acid hydrolysis of different chemically phosphorylated
raw starches have confirmed that phosphorylation favours
a restriction towards both types of hydrolysis (Sitohy &
Ramadan, 2001). However, for swollen or partially swollen
granules, phosphate groups repel each other and conse-
quently may promote amylase hydrolysis of starch granules
(Slaughter, Ellis, & Butterworth, 2001). No studies have
been reported on the effect of native phosphorus of starch
granules on the in vitro digestibility.
Food processing and starch digestibility
A wide range of techniques is being used by the industry
for processing various food materials. Processing leads to
an alteration in the food structure and also influences the
nutritional characteristics of the food including starch di-
gestibility. The outcome of some recent studies related to
different processing techniques and their effect on starch
digestibility is presented in Table 3. When starch molecules
are heated in excess water, the crystalline structure is dis-
rupted and water molecules become linked by hydrogen
bonding to the exposed hydroxyl groups of amylose and
amylopectin, which causes an increase in granule swelling
and solubility. Therefore, the water activity or the availabil-
ity of water is an important factor that determines the extent
of starch digestibility through enzymatic hydrolysis. The
Km of starch hydrolysis increased by a factor 2.5 e4 at
low moisture content compared to starch suspended in ex-
cess of water. It has also been observed that Vmax increased
with increasing moisture content of the hydrolysis reaction
(Slaughter, Ellis, Jackson, & Butterworth, 2002). Process-
ing has been observed to result in an increase in the degree
of starch hydrolysis, with values higher than 90% (at the
end of incubation with pancreatine) for wheat, barley and
oats (Anguita, Gasa, Martı́n-Orúe, & Pérez, 2006). Starches
in tubers and legumes are particularly well protected from
the polar environment of luminal fluids, and even cereals
such as wheat may not have access to a-amylase in the in-
testinal lumen unless they have been physically altered.
The principal process facilitating starch availability for
water penetration and consequent a-amylase action are
physical processing and cooking by heating to 100  C for
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J. Singh et al. / Trends in Food Science & Technology 21 (2010) 168e180 173

Table 3. Effect of processing on starch digestibility. formation of amylose lipid complexation, starcheprotein
interaction, and limited water availability which prolongs
Processing Starch Reference
digestibility
the starch digestibility during enzymatic hydrolysis
(Guha, Ali, & Bhattacharya, 1997). The decrease in the
Cooking 34a Roopa and Premavalli
(2008)
size distribution of the granule results in an increase in
Pressure cooking 42a the surface area. As a result, a process like grinding leads
Autoclaving 39.7a to a higher percentage of hydrolysis. Anguita et al.
Re-autoclaving 37.3a (2006) observed that extrusion provoked a decrease in par-
Puffing 33.4a ticle size compared to raw samples and affected the digest-
Roasting 37.2a
Baking 7.2a
ibility. Traditional and conventional processing methods
Frying 11.2a were compared with extrusion cooking and their effects
Germination 15.4a on bean starch digestibility were studied by Alonso et al.
Malting 15.5a (2000). Extrusion produced a higher increase in starch di-
Toasting 31.8a gestibility than other processing methods. Similarly, cook-
Gamma irradiation 75.2a Chung and Liu (2009)
Sheeting of pasta 156b Kim et al. (2008)
ing of legumes was carried out using boiling water and
dough (3 passes) with autoclaving at 121  C for up to 90 min and its effect
Sheeting of pasta 217b on starch digestibility was compared (Rehman & Shah,
dough (45 passes) 2005). Higher digestibility values were obtained which
Dehulled beans 151c Alonso et al. (2000) could be attributed the higher degree of starch gelatiniza-
Germinated (48 h) beans 178c
Extruded beans 306c
tion and destruction of anti-nutrients, when cooked by
Popped amaranth seeds 112d Capriles et al. (2008) autoclaving.
Cooked amaranth seeds 96d Processing of cereals such as dehulling, soaking and ger-
Flaked amaranth seeds 120d mination may result in an enhancement of digestibility due
Extruded amaranth seeds 93d to the loss of phytic acid, tannins and polyphenols which
a
Expressed as rapidly and slowly digestible starch (%). normally inhibit the activity of a-amylase and thus decrease
b
Expressed as rapidly and slowly digestible starch (%). the starch digestibility. It has been suggested that the re-
c
Expressed as starch digestibility (%).
d moval of tannins and phytic acid creates a large space
Expressed as hydrolysis index (%).
within the matrix, which increases the susceptibility
towards enzymatic attack and consequently improves the
starch digestibility (Rehman & Shah, 2005). An interesting
several minutes. Cooking increases the rate of hydrolysis by comparison of processing parameters such as popping,
gelatinizing the starch and making it more easily available roasting, flaking, and extrusion and their effects on ama-
for enzymatic attack. Bravo, Siddhuraju, & Saura-Calixto. ranth starch digestibility and predicted glycemic index
(1998) studied the effect of various processing methods has been reported recently in the literature (Capriles
such as direct cooking, with and without soaking and germi- et al., 2008). The authors reported that the rate of starch hy-
nation on the in vitro starch digestibility and resistant starch drolysis is significantly more enhanced by popping, roast-
content of Indian pulses. Sprouting and direct cooking re- ing and flaking than extrusion. Similarly, a range of
sulted in lowest resistant starch in freshly cooked and stored processing types was studied and their effects on in vitro di-
legumes, respectively while soaking increased the in vitro gestibility of finger millet starches were reported by Roopa
starch digestibility to a considerable extent. Processed le- and Premavalli (2008). The starch digestibility has been
gumes contain significant amounts of resistant starch in com- shown to increase by 35e40% during cooking, autoclaving
parison to other food products such as cereals and potatoes, and puffing followed by pressure cooking and germination.
irrespective of the processing treatment. The starch digestion They also reported that baking, frying and shallow frying
rate and the release of glucose in the blood stream are very reduced rapidly digestible starch (RDS) while roasting
low after the ingestion of processed legumes (Bravo et al., and pressure cooking enhanced the RDS to about 23% fol-
1998). lowed by cooking, autoclaving and puffing. The increase in
Extrusion cooking significantly increases the in vitro di- the sheeting cycles of pasta dough has also been observed
gestibility of starches (Alonso, Aguirre, & Marzo, 2000; to enhance the in vitro starch digestibility due to pulling
Altan, McCarthy, & Maskan, 2009). The increase in digest- away of protein from the starch granules (Kim et al.,
ibility of starch may be explained on the basis that the 2008). The reduction in cohesiveness between starch and
starch granules lose their structural integrity due to in- proteins of dough may increase the starch accessibility to
creased shearing action and kneading in the extruder barrel a-amylase which leads to greater digestibility. A detailed
which ultimately increase their susceptibility towards enzy- review has been published recently on the importance of
matic attack. Somewhat low values of digestibilities from structure development at microscopic and molecular level
the extrusion cooked starch or starchy foods have also during pasta making and its relationship with starch and
been seen sometimes which may be attributed to the protein digestibility (Petitot, Abecassis, & Micard, 2009).
 Author's personal copy
174 J. Singh et al. / Trends in Food Science & Technology 21 (2010) 168e180
The authors emphasized that the compact structure of pasta,
the encapsulation of starch by proteins and the physical
structure of starch are mainly responsible for the reduced
enzymic susceptibility of starch in cooked pasta.
Assimilation of starches may be enhanced by a series of
processes, such as cracking of the grain, and then converting
it from a crystalline to a gel structure, which promotes effi-
cient entry into the luminal polar solution for interaction
with the a-amylases (Goni et al., 1997). Effects of micro-
wave heating and other heating methods such as cooking
and pressure cooking on the rate of hydrolysis and glycemic
index were compared between kudzu starch and maize
starch (Geng, Zongdao, & Toledo, 2003). The rate of hydro-
lysis for both the starches increased following heat treat-
ment and to a greater extent with microwaving cooking,
which may be attributed to the greater penetration of heat
during microwave heating. The change in degree of suscep-
tibility of the starches towards enzymatic digestion is a func-
tion of the extent to which the microwave heating process
induced any changes in the crystalline structure of the
starches (Anderson & Goraya, 2006). Recently, the effects
of microwave treatment time on the digestibility of Canna
edulis starch have been reported by Zhang, Wang, & Shi
(2009). Their results indicated that both low (400 W) and
high (1000 W) microwave powers are advantageous to the
formation of RS. Irradiation has been used to extend the
shelf life and safety of food and is permitted now in many
countries. Degradation of starch polymers by gamma irradi-
ation may result in reduction in molecular weight of amy-
lose and amylopectin, decreased viscosity, and increased
acidity (Abu, Duodu, & Minnaar, 2006; Bao, Ao, & Jane,
2005). Chung and Liu (2009) studied the effect of gamma
irradiation on the enzymatic digestibility of corn starch
and reported that a slower dose rate decreases the RDS
and SDS contents and increases RS content.
During the gelatinization of starch, the crystalline struc-
ture of amylopectin disintegrates and the polysaccharide
chains take up a random configuration thus causing the
swelling and rupturing of the starch granules (Singh
et al., 2007). After gelatinization and cooling of the cooked
starch, re-crystallization of the starch chains starts to occur.
Amylose aggregation and crystallization in the cooked
starch pastes have been reported to be complete within
the first few hours while amylopectin aggregation and crys-
tallization occur at later stages during refrigerated storage
(Singh et al., 2006, 2008). The linear chains of amylose fa-
cilitate the cross-linkages through hydrogen bonds whereas
the branched amylopectin delays its recrystallization. The
retrogradation properties of starches are also influenced
by the structural arrangement of starch chains within the
amorphous and crystalline regions of the un-gelatinized
granule, because this structural arrangement influences
the extent of granule breakdown during gelatinisation and
also influences the interactions that occur between starch
chains during storage (Kaur et al., 2007). The resistant
starch formation in the cooked starches stored at
refrigeration temperatures is greatly influenced by the ex-
tent of retrogradation. Retrograded amylose from peas,
wheat and potatoes is highly resistant towards enzymatic
hydrolysis. It has been observed that storing cooked rice
at refrigerated temperatures may lead to a reduction in their
digestibility and estimated glycemic index (Frei et al.,
2003; Hu et al., 2004). The same phenomenon can be ob-
served in cooked starchy tubers. Potatoes were cooked
and then cooled at refrigeration temperatures for up to
2 days and their starch digestibilities were compared with
the freshly cooked ones by Mishra, Monro, & Hedderley
(2008). They reported that the percentage of rapidly digest-
ible starch in the refrigerated potatoes decreased to 45%
from 95% of the freshly cooked potatoes which could be
attributed to the retrogradation of starch and resistant starch
formation. The dispersed polymers of the gelatinized starch
during refrigerated storage have been observed to undergo
retrogradation which leads to the formation of to semi-crys-
talline structures that resist digestion by amylases.
Modified starch and starch digestibility
Native starch is a good texture stabilizer and regulator in
food systems, but limitations such as low shear resistance,
thermal resistance, and high tendency towards retrograda-
tion limit its use in some food applications. Chemical starch
modification is generally achieved through derivatization
such as etherification, esterification, cross-linking and
grafting of starch. Chemical modification involves the in-
troduction of functional groups into the starch molecule, re-
sulting in markedly altered functional properties. Such
modification of native granular starches profoundly alters
their gelatinization, pasting and retrogradation behaviour
(Singh et al., 2007). There are many reports available that
modifications of starch such as hydroxypropylation, octe-
nylsuccinylation, and combinations of cross-linking and
substitution reduce the extent of their enzyme-catalyzed hy-
drolysis (Han & BeMiller, 2007). After carrying out chem-
ical modifications on maize starches, Chung, Shin, & Lim
(2008) suggested that substitution and oxidation contributes
to increasing the resistant starch content of the starches
whereas cross-linking does not affect starch digestibility
to a considerable extent (Fig. 1). However, studies by
Hwang et al. (2009) on the in vitro digestibility of hydrox-
ypropylated and cross-linked waxy and non-waxy starches
have shown no significant changes in resistant starch con-
tent compared to control unmodified starch samples. It is
important to point out that the general resistant starch
assays may not be appropriate to measure the resistant
starch content of chemically modified starches. It has
been suggested that the pancreatinegravimetric method
involves exhaustive digestion and is therefore more appro-
priate for the determination of resistant starch in chemically
modified starches (Hwang et al., 2009). Many modified
starches made for food use contain only small amounts of
substituent groups and have been used as safe food ingredi-
ents. The legislative approval for the use of novel starch
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J. Singh et al. / Trends in Food Science & Technology 21 (2010) 168e180
derivatives in processed food formulations is still under de-
bate, but several tailor-made starch derivatives with multi-
ple modifications are being prepared and characterized
(Tharanthan, 2005). Some of the chemically modified
starches are being increasingly used as fat replacers or fat
substitutes in different food systems. These starches are
either partially or totally undigested, therefore contribute
zero calories to the food on consumption (Tharanthan,
2005). The chemical modification of starch by acetylation
also improves the satiating, glycemic and insulinemic prop-
erties of the meal (Raben, Andersen, Karberg, Holst, &
Astrup, 1997). Phosphorylated/cross-linked starches are
slowly digestible and are thought to provide nutritional
benefits for humans (Sang & Seib, 2006). Also the slowly
digesting modified starches could be used for the treatment
of certain medical modalities (e.g. glycogen storage disease
and diabetes mellitus) (Wolf, Bauer, & Fahey, 1999). Sev-
eral studies have highlighted various methods and technol-
ogies (other than chemical modification) to modify the
starch for the formation of SDS (Lehmann & Robin,
2007, and references therein). The prominent methods to
produce SDS include annealing and heat moisture treat-
ments of starch (Anderson et al., 2002; Lehmann et al.,
2002; Lehmann & Robin, 2007; Shin et al., 2005).
Viscosity of the food matrix and starch digestibility
Among the several factors that can affect the enzymatic
digestibility of starch and glycemic response, reports are
also available on the effects of the food matrix viscosity.
The physical texture of the food may affect the digestion
of starch and the absorption of hydrolysis products. Poly-
saccharide based gums belong to the water-soluble non-
starch polysaccharides and their effects on the human
metabolism are considered to be beneficial because they
Fig. 1. Starch hydrolysis patterns of the chemically modified maize
starches in the prime (granular) state with porcine pancreatic a-amy-
lase for 3 h at 37  C. PUM, unmodified starch in the prime state;
AC, acetylated starch; OX, oxidized starch; HP, hydroxypropylated
starch; and CL, cross-linked starch (reprinted from Chung et al.,
2008 with permission from Elsevier).
175
decrease postprandial glycaemia following ingestion of
starchy food (Kaur & Singh, 2009). They are used in
starch-based products as a processing ingredient (thicken-
ing agent, emulsifier, stabilizer). Ellis, Roberts, Low, &
Morgan (1995) studied the effect of adding guar gum (at
different concentrations) on the net apparent glucose ab-
sorption in growing pigs. The pig meals containing guar
gum resulted in increased zero shear viscosity of jejuna
digesta along with a significant reduction in the rate of glu-
cose absorption. This postprandial effect of guar gum re-
sults because of the gum’s capacity to increase the
viscosity of digesta within the gastrointestinal tract due to
the enlargement of fully hydrated galactomannan chains.
This whole phenomenon reduces the rate of digestion and
absorption of carbohydrates and therefore lowers the post-
prandial rise in blood glucose. The presence of galacto-
mannan in the starch mixture impose restrictions on the
swelling of starch granules during gelatinization which re-
sults in the size reduction of starch granule remnants in the
cooked starch paste whereas some of the granules may also
not gelatinize properly because of less availability of water
molecules to starch granules (Kaur, Singh, Singh, & Mc-
Carthy, 2008). This incomplete gelatinization of starch
granules may also increase their resistance towards enzy-
matic hydrolysis. The viscous fibre derived from different
gums such as guar, tragacanth increases the viscosity
even at relatively low polymer concentration in the food
matrix which may increase the overall viscosity of digesta
in the gastrointestinal tract. The consequence is the de-
creasing of the postprandial carbohydrate absorption after
ingestion of starchy food.
The increase in viscosity of digesta can affect gastric
function and inhibit propulsive and mixing effects gener-
ated by peristalsis. Under these conditions, interactions be-
tween substrates and digestives enzymes are less frequent.
Apart from low rate of starch digestion, the transport of am-
ylolytic products (e.g. maltose, a-limit dextrins) has also
been slowed down through mucosa, when gums are present
(Brennan, Blake, Ellis, & Schofield, 1996). The addition of
some cereal based viscous fibre in meals also influences the
digestion and absorption of carbohydrates to a considerable
extent. However, the postprandial glucose response may be
slightly different as they are physically and functionally
different from the viscous gums.
In addition, it may be difficult to distinguish among the
two effects whether decrease in the rate of hydrolysis occur
by the inhibition of enzymes or by an increase in the viscos-
ity of gastrointestinal contents (acting on the mass transfer).
An inhibitory action of very small concentration (0.5%) of
galactomannan addition on amylase acting on a range of
concentrations of gelatinised starch was observed by
Slaughter et al. (2002). The Km for the hydrolysis reaction
was not affected significantly by the presence of guar gum,
but kcat/Km decreased considerably. Such action would be
normally interpreted as a non-competitive effect on starch
hydrolysis.
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176 J. Singh et al. / Trends in Food Science & Technology 21 (2010) 168e180
Brennan et al. (1996) studied the addition of guar gum in
white bread and studied its microstructure along with in vitro
and in vivo digestibility. It was observed that the blood glu-
cose lowering action of gum is due to its ability to act as
a physical barrier to starch digestion along with increasing
the viscosity of digesta. The association between guar galac-
tomannan and starch has also been confirmed through the
microstructural analysis of pig digesta. However, according
to Slaughter et al. (2002), the existence of a significant phys-
ical barrier to the enzyme resulting from the galactoman-
nanestarch interaction is unlikely under the conditions
selected for their experiments (a concentration range of
guar galactomannan up to and including 1% by weight in
the final concentration) because of the lack of effect on Km
and a lack of time dependency of inhibition. It implies that
there are still sufficient unmasked attack sites on the granule
where amylase can act. Moreover gums contribute to de-
crease the water activity during hydrothermal processing of
the starch that could influence the a-amylase action and in-
fluence the starch swelling power. Chaisawang and Suphan-
tharika (2005) demonstrated that xanthan inhibited swelling
and prevented amylose leach out, whereas guar gum seemed
to have no effect on these properties. The ionic interactions
between starch and gum play an important role on the viscos-
ity and visco-elasticity characteristics. The strong electro-
static interactions between cationic starch and anionic gum
results in an instantaneous aggregation of granules, whereas
non-ionic gums form a sheet structure and loosely wrap the
granules (Chaisawang & Suphantharika, 2005). These obser-
vations are related to the changes in rheological parameters
such as storage modulus (G0 ) and loss modulus (G00 ) of
starchexanthan and starcheguar systems (Chaisawang &
Suphantharika, 2005; Kaur et al., 2007). However, to our
knowledge, no reports are available on the effect of ionic in-
teractions of the gums during in vitro starch hydrolysis.
Koh, Kasapis, Lim, & Foo (2009) studied the effect of the
addition of alginate on the in vitro digestion of rice dough
and reported a decrease in starch hydrolysis with the addi-
tion of sodium alginate. Alginate forms a continuous net-
work by suspending the starch granules in a coherent gel
which acts as a barrier when enzymes try to access the
starch. They further postulated that starch based alginate net-
works may help in the development of rice-dough based for-
mulations with reduced glycemic index. The presence of
non-starch polysaccharides from different plant sources
may affect the physical properties of the digesta at all sites
of the gastrointestinal tract. The high level of viscosity slows
down many of the physiological processes associated with
the digestion of foods and absorption of nutrients and thus
helps in improving the management of glucose intolerance
and obesity. The well-documented blood glucose reducing
effect of dietary supplements of water soluble non-starch
polysaccharides such as pectin and guar gum also depends
on their capacity to increase viscosity of digesta in the stom-
ach and the small intestine (Johansen, Knudsen, Sandström,
& Skjøth, 1996). Gastric response to increased meal
viscosity has been assessed by techniques such as echo-pla-
nar magnetic resonance imaging (Marciani et al., 2000).
Food proteins and starch digestibility
The presence of protein in the food matrix may influence
the rate of starch digestion. Digestibility of the starches and
proteins in various cereal products is significantly affected
by their interaction with each other. The functional proper-
ties and starch digestibility have been observed to be influ-
enced by the presence of even small amounts of protein in
cereals and other food products (Ezeogu, Duodu, Emman-
bux, & Taylor, 2008). Protein fractions such as albumin,
globulins and glutenins help in gluing the protein bodies
into a matrix surrounding starch granules that may act as
a barrier towards starch digestibility (Hamaker & Bugusu,
2003). Cooking or processing may sometimes reduce the
starch digestibility as the conformational changes in pro-
teins may occur that could facilitate the formation of disul-
fide-linked polymers (Oria, Hamaker, & Shull, 1995). The
presence of a protein barrier surrounding the starch granule
has been confirmed by the addition of pronase enzyme to
hydrolyze the protein matrix and a significant enhancement
of in vitro starch digestibility was observed afterwards due
to the clearance of passage for amylase and amyloglucosi-
dase (Rooney & Pflugfelder, 1986). Wong et al. (2009)
have reported that a greater abundance of disulfide-bonded
proteins, the presence of non-waxy starch and the granule
bound starch synthase enzyme may affect the digestibility
of both starch and protein in sorghum grain endosperm. An-
other study by Choi, Woo, Ko, & Moon (2008) reported an
increase in the in vitro starch digestibility when sodium sul-
phite was added during cooking of waxy sorghum flour.
The reducing agents such as sodium sulphite or bisulphite
may prevent the formation of enzyme resistant disul-
phide-linked plant polymers which facilitate easy access
of amylolytic enzymes to the starch granule. Effects of
the protein matrix on in vitro starch digestibility of pro-
cessed starch products such as pasta have also been re-
ported (Kim et al., 2008). Jenkins et al. (1987) studied
the effect of starcheprotein interaction in wheat and its ef-
fect on starch digestibility. Their reports suggested that the
occurrence of a starcheprotein interaction in white flour
may account for the decreased glycemic response and re-
duced rate of digestion. It has already been reported
(Liener, 1980) that 10e20% of the starch in white wheat
flour may be malabsorbed as judged by breath-hydrogen
production. Subsequent studies verified that significant
amounts of starch may indeed enter the colon (Wolever
et al., 1986). It has been observed that removing gluten
from wheat flour eliminated the starch malabsorption but
this effect was not reversed by subsequently adding back
the gluten to the gluten-free flour. This raised the question
of whether the natural starcheprotein interaction is respon-
sible for the reduced digestibility of starch (Jenkins et al.,
1987). In vitro digestion studies showed that the concentra-
tion of total starch-digestion products was significantly
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J. Singh et al. / Trends in Food Science & Technology 21 (2010) 168e180
lower for white bread than for gluten-free bread. In terms of
possible mechanism, all-purpose wheat flour is composed
of granules with a starch core surrounded by a protein net-
work. The protein network may inhibit the rate of hydroly-
sis in the lumen of the small intestine (Jenkins et al., 1987).
Food lipids and starch digestibility
The amylose chain of starch displays a natural twist pro-
viding a helical conformation with six anhydroglucose
units per turn (Zobel, 1988). Hydroxyl groups of glucose
residues are present on the outer surface of the helix, while
the internal cavity is a hydrophobic tube (Zhou, Robards,
Helliwell, & Blanchard, 2007). The hydrophobic complex-
ing agents can stay or complex within the amylose helix
stabilized through van der Waals forces with the adjacent
C-hydrogen of amylose (Godet, Tran, & Delagw, 1993;
Zhou et al., 2007). The effects of free fatty acids (lauric,
myristic, palmitic, stearic and oleic acids, lysolecithin and
cholesterol) on the hydrolysis of starch, amylose and amy-
lopectin using a-amylase and amyloglucosidase have been
reported in the literature by Crowe, Seligman, & Copeland
(2000). Around 60% amylose was converted to glucose in
1 h, reaching up to 90% after 6 h. The addition of lauric,
myristic, palmitic and oleic acids reduced the enzymatic
hydrolysis of amylose by 35%. However neither stearic
acid nor cholesterol presented an inhibition. Lauric acid
had no effect on the enzymatic breakdown of amylopectin,
whereas the breakdown of whole starch was inhibited 12%
by lauric acid. These experiments suggest that only the hy-
drolysis of the amylose fraction (31% of the whole starch)
is affected by lauric acid. Amylose presents a helical con-
formation and can form inclusion complexes with small hy-
drophobic molecules. Complexes between fatty acids such
as lauric acid and amylose can form rapidly under physio-
logical conditions which contribute to the formation of re-
sistant starch (Seligman, Copeland, Appels, & Morell,
1998). The formation of such complexes with lipids may
result in significant changes in the behaviour of the starch,
including decreased solubility, increased gelatinization
temperature and delayed retrogradation and resistance to-
wards the action of digestive enzymes. Amylose may
bind one lauric acid molecule per w20 glucose units in
the glucose chain, but in contrast, very little lauric acid
binds under the same conditions to amylopectin and other
branched glucans (Crowe et al., 2000).
Enzymatic resistance of the pure amylose and lipid com-
plexes has also been reported in the literature (Gelders,
Duyck, Goesaert, & Delcour, 2005; Holm et al., 1983).
With the help of in vitro and in vivo digestibility studies on
amyloseelipid complexes, Holm et al. (1983) observed
that complexed amylose is hydrolysed and absorbed in the
gastrointestinal tract to the same extent as free amylose but
at a somewhat slower rate. After studying the influence of en-
zymatic action on the digestibilities of complexes formed be-
tween amylose of different average chain lengths (degree of
polymerisation) and docosanoic acid/glycerol monostearate,
177
Gelders et al. (2005) suggested that enzymatic resistance of
complexes increases with increasing amylose degree of
polymerisation, lipid chain length and complexation temper-
ature. They further reported that enzymatic hydrolysis of
these complexes gives rise to two or more dextrin subpopu-
lations, which originates from a sequence of lamellar units
with interconnecting, amorphous amylose chains.
a-Amylase inhibitors, anti-nutrients and starch
digestibility
The more common types of a-amylases are of protein or
glycoprotein nature, non-competitive, non-dialyzable and
generally heat labile (Dreher et al., 1984). A wide variety
of food crops such as beans, rye, wheat and oats contain am-
ylase inhibitors in varying quantities. Non-proteinaceous am-
ylase inhibitors such as polyphenolic compounds or phenolic
acids, acarbose, isocarbose, acarviosine-glucose have also
been reported (Farias et al., 2007). Some oligosaccharides
derived from microbes also produce inhibitors that may in-
hibit the amylase activity. Amylostatin and acarbose are
two of the a-amylase inhibitors produced by a fungi Strepto-
myces and bacteria from family Actinoplanaceae, respec-
tively. Many molecules that exist in plant sources are
capable of inhibiting the activity of a-amylase. Results
from clinical studies have shown that the natural inhibitors
isolated from white beans significantly reduce the peak of
postprandial glucose in healthy and type 2 diabetic subjects
(Boivin, Flourie, Rizza, Go, & DiMagno, 1988). a-Amylase
inhibitors of wheat do not affect the wheat amylase activity
but are known to inhibit mammalian salivary and pancreatic
a-amylase (Lankisch, Layer, Rizza, & DiMagno, 1998).
Some low molecular weight plant-derived molecules like lu-
teolin, strawberry extracts, and green tea polyphenols have
also been observed to inhibit a-amylase or reduce postpran-
dial hyperglycemia (He et al., 2006; McDougall et al., 2005).
a-Amylase inhibitors from kidney beans have the highest
inhibitor activity although it is pH dependent. Some of these
inhibitors have a greater affinity for salivary amylases than
pancreatic amylases. The amylase inhibitors present in legume
crops are generally inactivated at or above 100  C. Amylase
inhibitors tend to be specific against particular amylases.
The inhibitor forms a complex with amylase and causes reduc-
tion in starch digestion. Many inhibit the activity of exogenous
a-amylases such as those produced by insects and may play
a role in protecting the seed against predation. Others affect en-
dogenous a-amylases and as such may participate in regulat-
ing a-amylase activity during seed development and
germination. Most amylase inhibitors from plants are active
against animal amylases. A natural amylase inhibitor has
been isolated from the germ fraction of wheat, which appeared
very labile. It is indeed destroyed by heating treatment and by
passage through the roller mill (Snow & O’Dea, 1981). Vari-
ous commercial starch blocking products have been manufac-
tured since early 1940s but many of the clinical studies have
proved that they do not work in vivo whereas some studies
have reported their positive role during in vitro experiments.
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178 J. Singh et al. / Trends in Food Science & Technology 21 (2010) 168e180
a-Amylase inhibitors have been found to be unstable in the
stomach and active only after pre-incubation with amylase
in the absence of starch (Lajolo and Genovese, 2002).
Dietary fibre has been suggested as the primary factor
influencing the slower rate of glucose release in foods
through its high viscosity which slows down gastric empty-
ing absorption of digested products in the small intestine.
The rate of starch digestion, however, cannot be explained
by the amount of fibre alone since some legumes (e.g., len-
tils) have similar fibre contents as certain cereal products
(e.g., wholemeal bread) and yet are digested at very differ-
ent rates in vitro (Jenkins et al., 1980). The high concentra-
tion of anti-nutrients such as phytic acid, lectins, enzyme
inhibitors in legumes may also play a role in starch digest-
ibility. Enzyme inhibitors and lectins have also been shown
to produce hypoglycemia and to decrease growth rates in
rats (Thorne et al., 1983).
Phytic acid is the most important phosphate reserve com-
pound in many plants. It can form a complex with proteins
and/or metal ions, reducing their biological availability.
Yoon, Thompson, & Jenkins (1983) have shown that the ad-
dition of phytic acid after pre-incubation with saliva de-
creased the sugar liberation significantly while little effect
has been seen on simultaneous addition. The phytic acid
may affect the starch digestibility through interaction with
amylase protein and/or binding with salivary minerals such
as calcium which is known to catalyze amylase activity.
The effects of processing on anti-nutrients (phytic acid, con-
densed tannins, polyphenolics, a-amylase inhibitor) acting
on starch digestibility are also important to consider (Alonso
et al., 2000; Rehman & Shah, 2005). For instance, the most
efficient treatment for phytic acid reduction found was ger-
mination for 72 h. Extrusion cooking also causes a significant
reduction in phytic acid and in condensed tannins and poly-
phenolics as well. Thermal processing methods have been re-
ported (Rehman & Shah, 2005) to act by thermal degradation
of the anti-nutrient molecules, changes in their chemical re-
activity and formation of insoluble complexes.
Conclusion and future research directions
Factors acting on starch digestibility are multiple and not
necessarily unrelated to one or more of the others. The impor-
tant factors that can be considered to have an influence on the
catalytic efficiency of the enzymes responsible during in vitro
starch hydrolysis are the starch characteristics, the physical
access of the enzyme to the starch, the availability of water
needed for the hydrolysis of the glycosidic linkage and the
lowered rates of diffusion of substrates due to the relatively
high viscosity. The effects of galactomannan such as guar
gum in lowering postprandial rise in blood glucose not only
include actions on intestine physiology but also include a di-
rect inhibition of the first step of the biochemical degradation
of starch. Other factors such as processing techniques, the
presence of other food components like proteins, lipids,
anti-nutrients/inhibitors also affect starch digestibility to
a significant extent. Based on the knowledge gained through
this review, the authors feel that more research should be di-
rected to explore the interactions between starch and non-
starch components of the food matrix and how in turn, these
interactions affect the digestibility of starch. Use of novel
plant based polysaccharides and new processing techniques
should be considered for the development of starch based
foods with low glycemic index.
Acknowledgements
The authors thank Professor Harjinder Singh, Co-Director
(Riddet Institute) for providing valuable comments and sug-
gestions on this manuscript.
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