Starch digestion capacity of poultry1
B. Svihus2
Norwegian University of Life Sciences, PO Box 5003, N-1432 Aas, Norway
ABSTRACT Starch is quantitatively the most impor-
tant nutrient in poultry diets and will to a large extent
be present as intact starch granules due to very limited
extent of gelatinization during pelleting. Although na-
tive starch is difficult to digest due to a semi-crystalline
structure, even fast-growing broiler chickens appears to
be able to digest this starch more or less completely
during passage through the jejunum. However, reduced
starch digestibility has been observed, particularly in
pelleted diets containing large quantities of wheat. Al-
though properties of the starch granule such as size and
Key words: amylase, broiler, enzyme
INTRODUCTION
Among the nutrients in poultry diets, starch is quan-
titatively the most important. Diets may contain up to
50% starch on a DM basis, and starch is the most im-
portant source of energy. This is a logical consequence
of cereals being the most important ingredient in poul-
try diets, and although Klasing (2005) has stressed that
galliformes are omnivorous and not granivorous as of-
ten stated, the capacity for digesting starch is high, as
will be discussed below. It is also well known that the
capacity for digesting fat is limited, at least in young
birds (Wiseman et al., 1998), and thus starch must be
the main energy source in diets. The other potential
energy sources of sugar and protein are, at least from
an economic and technical point of view, not suitable as
primary energy sources.
The main source of starch is seeds from plants in
the grass family, where starch is stored and used as
an energy source by the offspring during germination.
Although this source is predominantly in the form of
maize, wheat, and other cereals, starch from tubers
such as cassava may also be used in poultry diets.
©2014 Poultry Science Association Inc.
Received January 13, 2014.
Accepted May 10, 2014.
1 Presented as part of the Symposium: Possible Substrates for Ex-
ogenous Enzymes at the Poultry Science Association’s annual meeting
in San Diego, California, July 22 to 25, 2013.
2 Corresponding author: birger.svihus@nmbu.no
2394
components on the granule surface may affect digest-
ibility, the entrapment of starch granules in cell walls
and a protein matrix may be even more important fac-
tors impeding starch digestion. In that case, this and
the fact that amylase secretion is normally very high
in poultry may explain the lack of convincing effects
of exogenous α-amylase added to the diet. However,
few well-designed experiments assessing mechanisms of
starch digestion and the effect of α-amylase supplemen-
tation have been carried out, and thus more research is
needed in this important area.
2014 Poultry Science 93:2394–2399
http://dx.doi.org/10.3382/ps.2014-03905
STARCH STRUCTURE
Starch chemical structure and digestibility in gen-
eral (Svihus et al., 2005; Tester et al., 2006; Singh et
al., 2010), and for broiler chickens specifically (Carré,
2004; Svihus, 2011) have been reviewed previously. Two
distinct populations of starch exist. Amylopectin con-
sists of α-1–4 glucose chains with frequent branches
due to α-1–6 bonds, whereas amylose is characterized
by very few branches. Indeed, no structural continuum
is observed between these 2 types of α-glucans (Buléon
et al., 1998). Usually, less than half the amylose will
be branched and the number of branch points will be
less than 20 per molecule, whereas for amylopectin the
average number of branch points will be approximate-
ly once per 20 glucose units (Hizukuri et al., 1997).
Whereas amylose has a molecular weight of around 100
kDa, amylopectin has a much higher molecular weight
in the order of 104 to 106 kDa (Buléon et al., 1998).
Amylose forms double helices or single helices in the
native state (Buléon et al., 1998). Single helices give
rise to a central cavity that can be filled with com-
pounds such as iodine, alcohols, or fatty acids. Most
starches contain between 200 to 250 g of amylose/kg,
although some waxy starches contain very little, and
other starches, such as amylomaize, may contain 650 to
700 g of amylose/kg (Parker and Ring, 2001). In the na-
tive state, starch is organized in very complex and large
structures, where amorphous and crystalline layers al-
ternate to form rigid, semi-crystalline granules, varying
in size from 1 to 50 μm. Although the starch granule
 SYMPOSIUM: SUBSTRATES FOR EXOGENOUS ENZYMES
structure is far from completely understood (Perez and
Bertoft, 2010), the semi-crystalline layer is believed to
consist of alternating layers of crystalline α-glucans ex-
tending from intermittent branches of amylopectin and
amorphous amylopectin branch points, respectively. It
has been hypothesized that 1 growth ring is laid down
per day due to variation in photosynthetic activity and
thus access to glucose (Tester, 1997; Smith, 2001). As
discussed by, for example, Tester et al. (2004), starch
granule architecture will vary among cereal sources
both in regard to size and shape of the granules and to
the molecular architecture of the granule. Wheat, rye,
and barley have lenticular large (10–40 µm) and spheri-
cal small (2–10 µm) granules in a bimodal distribution,
whereas maize and sorghum have spherical/polyhedral
granules (2–30 µm) with a unimodal distribution.
STARCH DIGESTION
It is generally accepted that the highly organized
structure will pose a challenge for starch digestion,
and that native starch therefore will be incompletely
digested by many species. Björck et al. (2000) stated
that the degree of crystallinity is inversely related to
the rate of starch digestion, and Zhang et al. (2006a,b)
substantiated this by concluding that the slow digest-
ibility of native starch is related to the ordered struc-
ture of alternating crystalline and amorphous layers in
the starch granule. In fact, Zhang et al. (2006a) showed
that digestion of starch granules starts at surface pores
and interior channels, which allows for the amylase to
enter the interior and digest the granule gradually from
the inside.
When native starch is exposed to high temperatures
with water present, the granular structure will disinte-
grate in a process called gelatinization, and starch will
appear as an amorphous water-soluble mass. With an
excess water content (e.g., above 40%), gelatinization
temperature for most cereal starches ranges between
50 and 70°C. The gelatinization process renders starch
molecules more available for α-amylase, facilitating
starch digestion, although consistent improvements in
starch digestibility have not been observed for all spe-
cies [e.g., in broilers fed starch that has been completely
gelatinized in an extrusion process (Plavnik and Sklan,
1995; Zimonja and Svihus, 2009)]. Several investiga-
tions, using enzymatic or calorimetric methods, have
shown that due to limited moisture content and moder-
ate temperatures during conventional pelleting, only a
small amount of starch gelatinization, varying between
5 and 30%, occurs (Skoch et al., 1981, 1983; Goelema
et al., 1999; Svihus et al., 2004; Moritz et al., 2005;
Zimonja et al., 2007, 2008; Zimonja and Svihus, 2009).
Even under more severe temperature and shear treat-
ments such as expander-pelleting, starch gelatinization
will not be complete. Starch gelatinization of between
22 and 36% has been reported for expanded and pel-
leted feeds (Goelema et al., 1999; Cramer et al., 2003;
Zimonja et al., 2007). Results by Cramer et al. (2003)
2395
showed no improvements in starch digestibility with
the application of expander treatment followed by pel-
leting. Similar observations were reported by Plavnik
and Sklan (1995), where no change in apparent starch
digestibility in broilers fed expanded, compared with
unprocessed, feeds was observed. Zimonja and Svihus
(2009) likewise failed to detect any significant improve-
ments in starch digestibility for broiler chickens as a
consequence of pelleting, although a complete gelati-
nization due to extrusion improved starch digestibil-
ity of wheat diets exhibiting low digestibility. Carré et
al. (1991) observed an increased starch digestibility of
pea-based diets after pelleting, but this may have been
due to the grinding effect during the pelleting process,
which released starch trapped in the protein matrix,
cell walls, or both.
DIGESTIBILITY OF STARCH IN POULTRY
Starch in poultry diets is to a large extent present
in the form of native starch granules, and keeping in
mind the complex digestion process described above, a
low starch digestibility, at least for fast-growing broiler
chickens that consume large quantities of starch, would
be expected. However, a very high starch digestibili-
ty is commonly observed even in young broiler chick-
ens. Several studies have shown that chicks are rap-
idly adapting to starch digestion when fed at hatch,
as indicated by high activity levels of disaccharidase
(Mahagna and Nir, 1996) and α-amylase (Sklan and
Noy, 2000) 2 d after hatch. Accordingly, Zelenka and
Ceresnakova (2005) found that the total tract starch
digestibility coefficient already exceeded 0.96 at 3 d of
age for broiler chickens. The same authors also found
that starch digestibility decreased (P < 0.01) linearly
with increasing age in fast-growing broiler chickens
but not in slow-growing layer chickens. Thomas et
al. (2008) observed that starch digestibility in broiler
chickens dropped from 5 to 7 d of age, but was restored
to a normal high level at 14 d of age. Even when mea-
sured on material collected from the ileum of broiler
chickens, starch digestibility has often been observed
to be above 0.95 even for pelleted diets (Svihus, 2001;
Hetland et al., 2002, 2003; Svihus et al., 2004; Hetland
et al., 2007). This high capacity of poultry for digesting
starch is truly impressive, not the least in broiler chick-
ens, where pelleted diets and a high appetite results
in material passing through the digestive tract in less
than 5 h (Svihus et al., 2002, 2010). This means that in
less than 5 h, the starch granules must be released from
the surrounding protein and cell walls, become com-
pletely moistened followed by complete degradation by
a cascade of amylases, and finally the resulting glu-
cose must be absorbed. This is particularly impressive
because in vitro studies have shown that intact nor-
mal starch granules after a pretreatment imitating the
preintestinal human digestion process are incompletely
digested even after 4 h under conditions resembling the
small intestine (Shrestha et al., 2012). Also, in pigs an
2396 Svihus
ileal digestibility of ungelatinized starch below 0.9 is
often observed (Stein and Bohlke, 2007; Willamil et al.,
2012), despite a considerably longer retention time of 4
to 10 h in the small intestine (van Leeuwen and Jans-
man, 2007; Wilfart et al., 2007). The capacity of broiler
chickens to digest even unprocessed starchy ingredients
is further illustrated by the fact that even when whole
untreated cereals have been used in large quantities,
starch digestibility has been observed to be very high.
Svihus et al. (1997) found ileal starch digestibility to
be 0.98 in a mash diet containing 70% untreated whole
barley as the only cereal source, and Svihus and Het-
land (2001) found that 50% of the birds exhibited an
ileal starch digestibility above 0.94, even when given
a pelleted diet with 38.5% whole wheat. Even more
astonishing were the results by Hetland et al. (2002),
who observed an ileal starch digestibility of 0.98 in di-
ets with 44% whole wheat mixed with other pelleted
ingredients. The starch digestion process in such cases
cannot commence before grinding in the gizzard, and
because digestibility was based on analyses of contents
collected from the ileal segment between Meckel’s di-
verticulum and the ileo-cecal junction, this means that
the whole starch digestion and glucose absorption pro-
cess has taken place during the short retention time in
the duodenum and jejunum, estimated by Rougiere and
Carré (2010) to be around 1 h.
The very high capacity of chickens to digest even na-
tive starch compared with other species such as pigs,
rat, and humans may be due to a particularly high
secretion of amylolytic enzymes through the pancre-
atic juice of chickens. Lehrner and Malacinski (1975)
found that amylase constituted a much larger propor-
tion of the protein in the pancreas from chickens than
from other animals such as rabbits and rats. However,
no comparative studies have been found, thus mak-
ing comparisons difficult due to the large variations
in methods used for measuring activity, target organ
for activity (pancreas, pancreatic juice, or intestinal
content), and way of expressing activity (total activity
in segment, activity per gram of content, activity per
gram of DM, activity relative to a marker, and activity
per gram of protein in content). A particularly relevant
value to compare would be amylase concentration in
the intestinal content. Hedemann and Jensen (2004)
and Hedemann et al. (2006) reported an amylase activ-
ity of 50 to 100 U/mL in small intestinal contents of
newly weaned pigs, and Fang et al. (2012) found levels
in pig jejunum to gradually increase from 73 U/mL just
after weaning to 148 U/mL 2 mo after weaning. Ren et
al. (2012), using a similar method for activity measure-
ments, found the amylase activity to be more than 400
U/mL in the jejunum contents of broiler chickens. The
latter result seems to be in line with results obtained
by Osman (1982) if a jejunal chyme volume of 10 mL
is assumed (no chyme volume was reported), who re-
ported a total amylase activity of around 4,000 U in
jejunal contents. Kadhim et al. (2011) compared the
pancreatic and intestinal levels of α-amylase between
wild red jungle fowl and commercial broiler chickens,
and found considerably higher levels in the latter. For
duodenal and jejunal amylase, the levels were between
2 and 3 times higher per gram of contents in broiler
chickens than in wild jungle fowl. Osman (1982) found
high levels of both glucoamylase and α-amylase in the
intestinal tract of chickens, and concluded based on a
very high activity in the jejunum and a very low activ-
ity in the ileum that the jejunum was the main site of
starch digestibility. Noy and Sklan (1995) also found
that the amylase activity was reduced toward the il-
eum, but still detected significant activities here. These
findings corroborate those of Riesenfeld et al. (1980),
who showed that starch digestion and glucose absorp-
tion to a large extent was completed by the time the
contents had reached the ileum.
Despite the above indications of high starch digestion
capacity, starch digestibility values below 0.9 (measured
on an ileal or total tract level, or both) have also been
seen in a large number of experiments (Wiseman et al.,
2000; Maisonnier et al., 2001; Marron et al., 2001; Svi-
hus, 2001; Svihus and Hetland, 2001; Weurding et al.,
2001; Carré et al., 2002; Hetland et al., 2002; Carré et
al., 2005; Zimonja and Svihus, 2009). In several of the
reports, a considerable variation among cereal species,
among varieties within species and between individual
birds, has been observed. This indicates that factors
intrinsic to cereals and birds alike are affecting starch
digestibility.
FACTORS IMPEDING STARCH
DIGESTIBILITY
It is clear that properties of the starch ingested will
affect digestibility, because numerous experiments
have reported that glucose response and digestibility
of starch vary with starch source (Svihus et al., 2005).
However, there is still a lack of knowledge on the exact
causes for these variations in starch digestibility. Fac-
tors such as the ratio between amylose and amylopec-
tin, granule size and content and properties of proteins,
lipids, and phosphates on the surface of starch granules
were identified as potential causes for low starch digest-
ibility (Svihus et al., 2005; Tester et al., 2006; Singh et
al., 2010). Although the complexity of this issue and
the lack of experimental data preclude firm conclusions,
these reviews indicate that a small granule size may
explain the high digestibility of starch from oats and
rice, and that a large gluten matrix may contribute to
low digestibility of starch from wheat.
As pointed out before (Svihus, 2011), wheat appears
to be predominant in papers reporting low starch di-
gestibility. Also, wheat has been shown to result in con-
sistently lower starch digestibility when used at a high
inclusion rate and when compared with other cereals
such as barley and oats (Svihus, 2001; Zimonja and Svi-
hus, 2009). Addition of xylanase has sometimes been
 SYMPOSIUM: SUBSTRATES FOR EXOGENOUS ENZYMES
shown to improve starch digestibility of wheat diets [see
Svihus (2011) for an overview of literature], indicat-
ing that fibers may affect starch digestibility. Although
this is correlated with reduced viscosity (Murphy et
al., 2009), the rather moderate correlation coefficient
indicates that other effects of enzyme addition are also
contributing. It is possible that accessibility of the
starch in the wheat endosperm is an issue, and that one
potential beneficial effect of enzyme addition is that
enzymes degrade cell walls and thus increase access to
starch and other nutrients in the endosperm cells, as
discussed by Murphy et al. (2009). This hypothesis is
supported by results by Amerah et al. (2009), where
xylanase addition improved ME content in hard wheat,
but not in soft wheat. Carré et al. (2005) found that
low starch digestibility was associated with hardness of
wheat, and investigated this further to elucidate causes
for low starch digestibility in hard wheat varieties. On
the basis of particle size analysis and microscopy of ileal
contents, they found that a large part of the undigested
starch was entrapped in cell wall material, particularly
from areas of the endosperm close to the aleurone layer
(Péron et al., 2007). The same authors did not find
a large number of particles in the size class of starch
granules in the ileum, which indicated that the low di-
gestibility observed with hard wheat was not caused
by structural arrangements of the starch granules. In
another experiment, it was shown that very fine grind-
ing corrected the very low starch digestibility observed
with a normal particle size distribution (Péron et al.
2005). As stated by Carré et al. (2007), this supports
a conclusion that the cause for a low digestibility of
starch in wheat diets is partly that starch granules are
entrapped in cell walls, protein matrix, or both.
It is possible that for broilers, the short time avail-
able for digestion may be one of the causes for im-
paired starch digestion under some circumstances. Re-
sults have shown that digestibility of a diet with a low
digestibility increases when feed intake is reduced by
changing diet form from pellets to mash (Svihus and
Hetland, 2001), and this indicates that feed intake may
be inversely correlated with starch digestibility. Several
studies have shown a significant negative correlation
between feed intake for individual birds on identical di-
ets and starch digestibility or AME value (Svihus, 2006,
2011). In data from one of these experiments, AME and
total tract starch digestibility for individual birds were
very strongly correlated (r = 0.984), and starch digest-
ibility was inversely related to feed intake as shown in
Figure 1 (Svihus, 2011). In these data, 4 out of 10 ad
libitum-fed birds on a finely ground pelleted wheat diet
showed signs of being feed overconsumers, character-
ized by a normal weight gain, a higher than average
feed intake, and an AME value <2,462 kcal/kg (Svihus
et al., 2010). The hypothesis that feed overconsumption
leads to an overly fast feed passage that results in poor
starch digestibility is consistent with observations by
Hughes (2008), who showed that AME of broiler chick-
ens increased with increasing transit time.
2397
Figure 1. Relationship between feed intake and starch digestibility
in individual broilers fed a wheat-based diet (from Svihus, 2011).
EFFECT OF SUPPLEMENTAL α-AMYLASE
Published work assessing the extent to which exog-
enous α-amylase may improve starch digestibility is
scarce. Mahagna et al. (1995), Ritz et al. (1995), and
Shapiro and Nir (1995) did not observe any improve-
ment in starch digestibility when α-amylase was supple-
mented to broiler chickens during the first 14 d of age,
and Moran (1982) stated that “unlike most mammals,
the ability of fowl to release sufficient amylase is never
a problem.” This corresponds with observations by Svi-
hus and Hetland (2001), where digestibility was not
improved by adding pancreatin to the water of broiler
chickens exhibiting low starch digestibility of a wheat
diet. Conversely, Gracia et al. (2003) observed a sig-
nificant increase in starch digestibility when α-amylase
was added to a maize-based diet, thus indicating that
α-amylase secretion may be a limiting factor. Jiang
et al. (2008) added increasing levels of α-amylase to
broiler chicken diets and observed an increase in weight
gain at the highest supplementation level. Interest-
ingly, endogenous α-amylase production seemed to be
reduced at this high level. Corroborating this, other
experiments in which amylase has been included in the
enzyme cocktail have shown improved nutrient utiliza-
tion (Cowieson et al., 2006; Cowieson and Ravindran,
2008; Olukosi et al., 2008), although because other en-
zymes were used together with amylase, these latter re-
ports are of limited value in this context. Interestingly,
Hughes et al. (1994) separated chicks from 2 breeds
based on genetic variants of the pancreatic α-amylase
followed by a growth trial, and concluded that one of
the α-amylase genotypes from one of the breeds re-
sulted in higher feed/gain, thus indicating that birds
of certain α-amylase genotypes may cause suboptimal
starch digestibility. If an impaired starch digestibility is
dependent both on specific properties of the diet such
as cereal type and inclusion level, and on bird-related
factors such as appetite and digestive tract develop-
ment, this may explain the conflicting and inconclusive
results. However, the very high starch digestibility ob-
served even under the most demanding conditions in-
dicate that broiler chickens have the capacity to digest
2398 Svihus
starch completely, and thus that Moran’s (1982) state-
ment that the bird releases sufficient amylase in most
cases may still be true.
The very few reports published on the addition of
α-amylase to poultry diets warrant further research
into this very important area of poultry nutrition. Ex-
periments should be carried out with pure α-amylases
tested with pelleted diets based on both wheat and
maize, and in different bird phases, because we have
seen that feed intake can play an important role. De-
tailed assessment of starch digestibility throughout the
small intestine should be also carried out. Furthermore,
experiments are warranted that address the causes of
effective starch digestion in broiler chickens, including
the effectiveness of intestinal chyme from broiler chick-
ens compared with other animal species in releasing
glucose from starch.
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