Livestock Science 100 (2006) 44 – 52

www.elsevier.com/locate/livsci

Starch digestion in horses: The impact of feed processing

V. Julliand a,*, A. De Fombelle b, M. Varloud b
a
Nutrition et santé digestive des herbivores, Enesad, B.P. 87999, 21079 Dijon Cedex, France
b
Evialis, 56250 Saint Nolff, France

Abstract

Neither the partition of digestion between the foregut and the hindgut nor the factors of variation for diverse starches in feeds
or rations is well documented in spite of their importance in respect of nutrition and health of the athletic horse. At similar
intakes, feed processing and the botanical origin of the starch are two major factors that control the extent of prececal starch
digestion. Physical and biochemical changes occurring during the process influence both the mean retention time of the feeds
and the enzymatic activity in the foregut. Apparent digestibility of cereal starch varies from 20% to 90% in the foregut
depending on the process used. Physical processes have a lesser effect than thermal and hydrothermal ones. Physical processes
increase significantly the prececal digestibility of cornstarch but have a moderate impact on other cereals. Starch digestibility is
increased by thermal and hydrothermal processes whatever the botanical origin. Feed processing was shown to affect the
fermentability of starch in ruminants. In horses, a similar impact is expected not only in the hindgut but also in the stomach
where numerous starch-utilizing bacteria have been observed. Further investigations are needed to identify the process which
allows the highest prececal digestibility and decreases the hindgut fermentability of starch.
D 2005 Published by Elsevier B.V.

Keywords: Starch digestion; Horses; Food processing

1. Introduction horses. Starch digestion varies depending on several

factors, the most important being the level of intake,
To meet their nutritional requirements, athletic the botanical origin, the genotype and the feed
horses are fed rich energy-dense concentrates that processing of the cereal grains.
contain a significant amount of f starch. The starch in Grains can be subjected to mechanical (rolling,
cereal grains is primarily composed of amylose and crushing, grinding), thermal (roasting, micronizing) or
amylopectin molecules arranged around a central thermo-mechanical (flaking, popping, extruding) pro-
hilum, and represent the major sources of starch, cesses into dry or wet (steam rolling, steam crushing,
e.g. oats, barley and corn that are fed to stabled steam flaking, pressure toasting, steam extruding,
pelleting) conditions. Although horse owners often
* Corresponding author. Tel.: +33 80 77 25 59; fax: +33 80 77 25
roll cereal grains, the most frequent processes used are
84. grinding, micronizing, flaking and pelleting. Whatev-
E-mail address: [email protected] (V. Julliand). er the process, the objective is to increase the
0301-6226/$ - see front matter D 2005 Published by Elsevier B.V.
doi:10.1016/j.livprodsci.2005.11.001
 V. Julliand et al. / Livestock Science 100 (2006) 44–52 45

nutritional value of the feeds by enhancing the digestive process, and results from these experiments
availability of starch. must be considered cautiously as they differ according
Researchers compared processing methods of to the time of collection after feeding (Kienzle, 1994).
grains and their effect on apparent digestibility of Cannulation of ponies and horses in the jejunum
starch in the total gastrointestinal tract (GIT) and (Gerhards et al., 1991) or ileum (Roger, 1989 in
indicated no significant change. In several different France; Peloso et al., 1994 in USA; Coleman et al.,
experiments, testing various feeding practices, starch 1988 in Canada and Leão et al., 1999 in Brazil) allows
digestibility was very high whatever the process: collection of samples that are representative of the
98.8 F 0.08%, 96.8 F 0.27%, 97.8 F 0.29% and prececal content (Kienzle, 1994) and can be informa-
99.5 F 0.13% on average for corn (Hintz et al., tive about the dynamics of the digestive process.
1971; Arnold, 1982; Wolter et al., 1982; Hinckle et When used on caecally fistulated horses or ponies,
al., 1983), oats (Householder, 1978; Arnold, 1982; the mobile bag technique (Macheboeuf et al., 1995;
Wolter et al., 1982; Hinckle et al., 1983; Brown, Moore-Colyer et al., 1997) is particularly successful
1987), barley (Arnold, 1982; McLean et al., for evaluating the disappearance dynamics of feed
1999a,b,c,d) and wheat (Wolter et al., 1982) respec- over time in the prececal segment of the GIT.
tively. However, an improvement in growth (Caine, The extent of digestion in the prececal segment is
1931 cited by Coleman, 2001) was reported suggest- determined by the enzymatic activity and by the
ing a better utilization of processed grains. Differences retention time of digesta in the foregut. Thus the
appear when partitioning starch digestibility. influence of grain processing on both the transit time
Depending on where starch is hydrolysed in the and the availability of foregut enzymes needs to be
GIT, the energetic efficiency of starch is highly identified.
variable: starch is transformed into glucose by the
host enzymes in the prececal compartments, whereas 2.1. Effect of the process on retention time and
it is degraded by microbial activity into volatile fatty enzymatic activity
acids (VFA) and lactate in hindgut fermentation
chambers. The partition of digestion between the 2.1.1. Retention time
foregut and the hindgut as well as the factors of In a recent review, Medina et al. (2002) reported
variation for diverse starches of the feeds or rations is that the mean retention time (MRT) of digesta in the
little documented despite their importance in terms of prececal part of the GIT is on average 6.8 F 1.2 h and
nutrition and health for the performance horse. At represents from 10% to 20% of the total GIT MRT. It
similar intakes and for a given cereal, feed processing varies from 1.6 to a maximum of 9.9 h, depending on
is expected to be a major factor controlling the extent many factors, including feed processing (Medina et
of prececal starch digestion. al., 2002). The MRT was reported to increase when a
The present review will discuss the impact of feed mixed diet was fed in a pelleted form compared with
processing on the prececal digestion and hindgut feeding the crude ingredients (Cabrera, 1995). The
fermentation of starch. MRT of the liquid phase and small particles (b 2 mm)
is shorter in the foregut than that of the solid phase in
most experiments (2 h versus 6 h on average) (Medina
2. The effect of processing on the prececal digestion et al., 2002).
of starch Although horses are known to masticate their food
into fine particles, grinding cereal grains reduces the
To evaluate the prececal digestibility of starch, particle size which affects their rate of passage.
three different techniques have been used on animals: Kleffken in 1993 (cited by Kienzle et al., 1997)
the collection from intestinal fistulated horses, the postulated that the soluble starch, liberated by
mobile bag technique and the postmortem collection extruded corn, could flow with the liquid phase and
of digestive contents. Postmortem collection of the thus have a shorter MRT than the insoluble form.
GIT contents is a drastic method that gives limited These observations emphasized the major role that
information in terms of explaining the dynamics of the processing can have on the transit of digesta.
46 V. Julliand et al. / Livestock Science 100 (2006) 44–52

Prececal retention time measures the total transit A particularly low amylytic activity was measured
between the mouth and the distal ileum without in the foregut of horses (Alexander and Hickson,
setting apart the different segments. Gastric emptying 1970): the a-amylase activity in the jejunum was
is part of the prececal transit and probably contributes estimated between 10 and 50 U/g (Meyer et al., 1993).
to its regulation. The rate and extent of the retention in When measured in the pancreatic tissue, this activity
the stomach has been recently evaluated with radio- was greater: from 85 to 909 U/g of tissue (Kienzle et al.,
labelled meal scintigraphy. The time necessary to 1994) but was still lower than that in other species, like
reach 50% of the remaining digesta (T50) of the solid the pig (3500 U/g in average) (Roberts, 1974; Frape,
phase is variable (from half an hour to 5 h) depending 1986). On the contrary, the activity of maltases and a-
on a number of factors. In particular, T50 is longer for glucosidases secreted by the equine intestinal mucosa
the solid phase than for the liquid one (Merrit, 1999), was very similar to that of other domestic mammals
and gastric emptying was shown to be influenced by (Frape, 1986). The secretion of amylase varies with the
meal size as well as meal composition (Roussel, diet, being maximal with an oatmeal compared to a hay
personal communication). meal and was reported to be independent of the grain
Little information is available about the transit of process (Radicke et al., 1992). The differences that
digesta in the small intestine. It is undoubtedly very were observed in the amylase efficiency can be
short which allows little time for the hydrolysation of explained by the composition of starch granules and
starch by the small intestine enzymes. The effect of the structure of the cell walls and the protein all of
the retention time on prececal starch digestibility is which influence the accessibility of the starch. Scan-
controversial: while McLean et al. (1999a,b,c,d) ning electron microscopic examinations showed that
reported an improvement of digestibility with longer the corrosion characteristics of the starch granules by
retention time, de Fombelle et al. (2001a) found no amylase in the equid foregut were typical for grains.
interaction between time and digestibility. Endocorrosion through pin holes was observed in
granules of corn, whereas on oats exocorrosion all
2.1.2. Enzymatic activity around the surface was noted, while barley starch
Due to a low a-amylase activity in saliva granules presented sporadic pin holes and small round
(Alexander and Hickson, 1970; Roberts, 1974; structures on the surface (Kienzle et al., 1997).
Radicke et al., 1992), oral starch disappearance is Interestingly starch degradation seems to be
probably negligible in the equid. In the stomach, independent of the amylase source: in vitro, starch
very high values of starch digestibility were reported: digestion was significantly modified by the various
50% (Varloud, personal communication), 58%, 69% types of starch but did not differ with the use of
(Wolter et al., 1980) and 75% (Wolter and Chaa- pancreatic amylase from different animal species
bouni, 1979). Recently de Fombelle et al. (2003) (Kienzle, 1988).
measured gastric starch disappearance from 95.9% to
97.9%, 65.6% to 70.7% and 23.9% to 27.2% for 2.2. Effect of the process on foregut digestion of starch
oats, barley and corn respectively. Because of the
weak activity of the saliva amylase, it was hypoth- Despite the importance of the stomach, the effect
esized that the abundant gastric microflora (Kern et of grain processing on the disappearance of starch in
al., 1974; Kienzle et al., 1997; de Fombelle et al., the gastric contents is poorly investigated. A compi-
2003) could contribute to the gastric disappearance lation of the current data suggests that expanding or
of starch in horses. The gastric pH of equines which pelleting significantly improves starch disappearance
is highly variable (from 1.5 to 7.0) (Merrit, 1999) in the stomach compared to grinding (Hintz et al.,
and can provide favourable conditions for microor- 1971; Wolter and Chaabouni, 1979; Wolter et al.,
ganism development. The activity of the microflora 1980; Varloud, personal communication). Also, feed
was demonstrated by the gastric concentrations of processing affects the fermentability of starch and this
lactate and VFA which are end products of the could have an impact on the gastric microflora
microbial degradation of starch (de Fombelle et al., altering the environment, and eventually causing
2003). conditions such as ulcers (Nadeau et al., 2000).
 V. Julliand et al. / Livestock Science 100 (2006) 44–52 47

100
At similar levels of starch intake, the extent of

Starch digestibility (% DM)

90
starch prececal digestion largely depends on its
80
botanical origin. Oat starch fed as a native form whole
70
showed the highest prececal digestibility (Household- crushed
60 ground
er, 1978; Kienzle et al., 1992). The digestibility of
50 popped
starch from corn grain was limited, due to the
40 pelleted
relationship with the protein matrix that encapsulates
30
starch granules and the compact nature of the starch
20
itself. According to these last observations, it appears 0 100 200 300 400
that the effect of grain processing on starch prececal Starch intake (g / 100 kg BW)
digestibility has to be observed for each cereal grain. Fig. 2. Prececal digestibility of cornstarch (Hintz et al., 1971;
Potter et al. (1992) suggested an upper limit to Arnold, 1982; Hinckle et al., 1983; Radicke et al., 1991; Meyer et
prececal starch digestion of 3.5 to 4 g starch/kg body al., 1993, 1995; de Fombelle et al., 2001a).
weight/meal of ground corn. More data are needed to
confirm the starch degrading capacity of the horse’s 1995; de Fombelle et al., 2003; Varloud, personal
foregut with ground corn and other grains before firm communication) (Fig. 3) and corn (Radicke et al.,
recommendations can be made. The intake level is 1991; de Fombelle et al., 2003).
another factor likely to mask the effect of grain Mechanical processing, and especially grinding,
processing. alters the physical form of grain primarily by breaking
Even if some results look contradictory, grain down the macrostructure of connected starch granules
processing generally improves prececal starch diges- and secondarily by destroying the structure of the
tion. The effect varies with the nature of the process starch granules themselves (Kienzle et al., 1997). The
used: amongst mechanical processes, rolling or surface area is thus increased which allows a greater
crushing did not significantly increase starch digestion exposure of the granules to digestive and microbial
in oats (Householder, 1978; Meyer et al., 1993, 1995) enzymes and enhances the rate of starch digestion.
(Fig. 1) and corn (Meyer et al., 1993, 1995) (Fig. 2) Probably rolling and crushing do not provide a finer
whereas grinding improved it for both oats (Radicke grinding than chewing the whole grain which explains
et al., 1991) (Fig. 1) and corn (Radicke et al., 1991; the lack of beneficial effect on starch digestion
Meyer et al., 1995) (Fig. 2) compared to the (Kienzle, 1994). Regarding grinding, the impact
unprocessed form of these cereals. In addition, no depends on the structure of the particles obtained
difference was noticed between rolling and crushing after the process. Under microscopic examination, it
but grinding significantly increased the prececal starch was observed that the ground corn with the lowest
digestibility of oats (Radicke et al., 1991; de Fombelle prececal digestibility showed very few single starch
et al., 2003), barley (Arnold, 1982; Meyer et al., 1993, granules and none were destroyed whereas that with

100 100
Starch digestibility (% DM)
Starch digestibility (% DM)

90 90
80 80
70 70 rolled

60 60 ground
whole
50 pelleted
50 rolled
40 ground 40
30 micronised 30

20 20
0 100 200 300 400 500 0 100 200 300 400
Starch intake (g / 100 kg BW) Starch intake (g / 100 kg BW)

Fig. 1. Prececal digestibility of oat starch (Householder, 1978; Fig. 3. Prececal digestibility of barley starch (Arnold, 1982; Meyer
Arnold, 1982; Radicke et al., 1991; Meyer et al., 1993, 1995; de et al., 1993, 1995; de Fombelle et al., 2001a; Varloud, personal
Fombelle et al., 2001a). communication).
48 V. Julliand et al. / Livestock Science 100 (2006) 44–52

Table 1 processing corn grain as they observed an increase of

Starch fractions of whole and extruded cereal grains and potato the glycemic response, used as an indirect measure of
starch at high (HT) or low (LT) temperature extrusion (from Murray
et al., 2001)
prececal starch digestibility.
It has been observed that the three starch fractions
% RDSa % SDSb % RSc TSd
[rapidly digestible starch (RDS), slowly digestible
Barley 23.2 11.4 17.0 51.6
starch (SDS) and resistant starch (RS) determined
LTe barley 30.3 13.5 4.8 48.6
HTf barley 47.8 4.4 6.0 58.2 using an enzymatic technique] of cereal grains and
Corn 34.6 14.6 23.6 72.8 potato starch were altered by heating (Table 1):
LT corn 54.2 13.1 6.4 73.7 extrusion increased systematically the concentration
HT corn 65.0 7.8 1.4 74.2 of RDS and substrates extruded at high temperature
Wheat 15.5 33.6 13.0 62.1
contained considerably higher concentrations of RDS
LT wheat 54.6 6.0 6.1 66.7
HT wheat 65.5 5.2 0.6 71.3 than those extruded at low temperature. On the
Sorghum 27.3 13.0 33.8 74.1 contrary, SDS and RS fractions decreased substantial-
LT sorghum 49.1 10.9 15.4 75.4 ly when substrates were extruded at high temperature
HT sorghum 70.0 5.4 2.1 77.5 compared to their native forms (Murray et al., 2001).
Potato starch 24.4 2.6 60.0 86.9
Heating modifies the structure of the grain: the
LT potato starch 65.4 27.0 2.2 94.6
HT potato starch – – – – grain macrostructure as well as the structure of the
a starch granule are destroyed and the biochemical
Rapidly digestible starch.
b
Slowly digestible starch. structure of starch is modified to a limited extent
c
Resistant starch. (Kienzle, 1994). Biochemical alterations of starch
d
Total starch = RDS + SDS + RS. increase with temperature, duration, water and me-
e
Extruded starch at low temperature (barley and corn, 83 8C; chanical processes. In the presence of water, heating
wheat, 86 8C; sorghum, 84 8C; potato, 94 8C).
f leads to the irreversible disruption of the crystallinity
Extruded starch at high temperature (barley, 135 8C; corn, 140
8C; wheat, 143 8C; sorghum, 145 8C). of the starch granule called gelatinisation of starch: the
greater the amount of water present and available to
the starch granule, the greater the extent of gelatinisa-
the highest digestibility presented many single par- tion. Highly gelatinised starch granules are often
ticles with some completely destroyed (Kienzle et al., shredded or distorted during processing. Gelatinisa-
1992; Meyer et al., 1993). Micronization was reported tion increases the solubility of starch and improves its
to enhance the digestibility of oat and sorghum starch susceptibility to enzyme attack.
compared to rolling (Householder, 1978). Thermo- The alterations of the starch granules after process-
mechanical processes significantly increased the ing was shown to correspond very well to starch
prececal digestibility of grain starch and popping digestibility in the jejunum and the difference in
and pelleting greatly improved cornstarch utilization digestibility between whole, crushed and ground corn
(Hintz et al., 1971; Meyer et al., 1993, 1995) (Fig. 2). clearly confirmed it (Table 2) (Kienzle et al., 1997).
Hoekstra et al. (1999) confirmed the positive effect of When popped corn was examined under a micro-

Table 2
Effect of processing corn on the botanical starch structure in the feed and pre-ileal starch digestibility in horses (Kienzle et al., 1997)
Feeds Apparent Connection of starch granules Structure of granules
digestibility (%)
Whole or crushed corn 29 Granules tightly fused, fitting together mosaic Similar size, middle sized (10–20 Am),
like polyhedric form, smooth surface
Ground corn, modern mill 47 Particles consisting of several connected Granules more or less unchanged
granules, all particles of similar size
Ground corn, farm mill 71 Many single granules but also larger particles Some granules damaged or destroyed
consisting of several granules
Expanded corn 90 Connection of granules partly or completely Granules deformed like melted, size
destroyed unaltered, partly amorphous structures
 V. Julliand et al. / Livestock Science 100 (2006) 44–52 49

scope, a complete destruction of the starch granules intake should not exceed 200 g starch/100 kg BW/
was observed or at least a considerable alteration was meal as suggested earlier (Kienzle et al., 1992; Meyer
seen (Meyer et al., 1995). In addition, the process et al., 1995). Medina (2003) compared apparent
interferes with other components of the grain that digestibility and cell-wall digestibility data from
probably also modifies the starch digestibility. different trials and concluded that a low proportion
of cereals, i.e. below 30% DM of the diet does not
interfere with the apparent digestibility of fibre and
3. The effect of starch processing on the hindgut indeed can improve it as it stimulates fibrolytic
fermentation activity. On the contrary, a larger proportion of cereals
in the ration negatively affects fibre degradation.
Whatever the grain and the process, starch digestion Current data do not permit conclusions about the
in the large intestine is almost complete: 91.1 F 6.8% effect of either mechanical (rolling, grinding) or
(Householder, 1978; Arnold, 1982; Hinckle et al., thermal (micronizing and pelleting) processes on the
1983; Brown, 1987). Despite this homogenous value, quantity of starch reaching the hindgut (Fig. 4).
some variations may appear amongst the microbial Starch undigested in the prececal segment is
population and the intestinal environment depending fermented by the microflora in the hindgut. The risk
on the quantity and the fermentability of starch flowing of inducing a dysfunction in the caecum or the colon is
into the hindgut ecosystem. higher if the grain is highly fermentable and if a high
A compilation of data on ileal starch digestion proportion of it arrives in the fermentation chambers
(Hintz et al., 1971; Householder, 1978; Arnold, 1982; (Kienzle, 1994). Although both cornstarch and barley
Hinckle et al., 1983; Brown, 1987; de Fombelle et al., starch are particularly resistant to prececal enzymatic
2001a; Varloud, personal communication) showed hydrolysis: barley is more fermentable in the hindgut
that from 4% to 30% of the ingested grain starch than corn (Kienzle, 1994). It seems that grain
can escape the prececal digestion (Fig. 4). When the processing could affect the extent to which starch is
level of starch intake was lower than 200 g/100 kg fermented by microorganisms: the starch of barley was
BW/meal, the prececally undigested starch never more degraded in the caecum when micronized or
exceeded 50 g/100 kg BW/meal. Above the limit of extruded than when rolled (McLean et al., 2000).
200 g/100 kg BW/meal, the quantity of starch Similarly, both micronizing and extrusion increased
reaching the hindgut fermenters varied from 70 to the degradability of starch in corn grain (McLean et al.,
150 g/100 kg BW/meal. This could confirm that the 1999b). This would be consistent with the increase in
the rapidly digestible starch fraction in cereal grains
that is caused by heating (Table 1) (Murray et al.,
160
2001). However, these data need to be interpreted
140
precaecally undigested starch

whole O cautiously as the measurements were conducted using

120 rolled O
whole cereal grains and may not be representative of
(g / 100 kg BW)

micronised O
100 pelleted O the degradability of prececally digested substrates.
80 rolled C Several authors have reported that disturbances due
60
ground C to the fermentation of starch in the hindgut can lead to
pelleted C
40
changes in the microbial profiles and activities as well
rolled B
pelleted B
as the environmental parameters (Radicke et al., 1991;
20
McLean et al., 1999a,b,c,d; Brown et al., 2001; de
0 Fombelle et al., 2001b; Julliand et al., 2001) which
0 100 200 300 400
Starch intake (g / 100 kg BW) can lead gut pathologies: Garner et al. (1978)
experimentally induced laminitis within 16 to 24
Fig. 4. Prececally undigested starch (g/100 kg BW) according to the h after ingestion of a ration containing 85% carbohy-
grain process for oats (O—round), corn (C—triangle) and barley
(B—square) (Hintz et al., 1971; Householder, 1978; Arnold, 1982;
drates. McLean et al. (2000) suggested that thermal
Hinckle et al., 1983; Brown, 1987; de Fombelle et al., 2001a; processes could be beneficial in minimising hindgut
Varloud personal communication). dysfunction: when ponies were offered 50:50 diets of
50 V. Julliand et al. / Livestock Science 100 (2006) 44–52

micronized or extruded barley and hay cubes the intra- Brown, W.Y., Roberts, K., Bird, S.H., Rowe, J.B., 2001. Safer grain
cecal fermentation parameters were largely similar to feeding for horses. Equine Nutrition and Physiology Sympo-
sium, (ENPS). Equine Nutrition and Physiology Society,
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in the literature. Moreover there is a paucity of Phillipeau, C., Jacotot, E., Julliand, V., 2001a. Effect of the
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