2006 Poultry Science Association, Inc.

Anatomy, Microbes, and Fiber:

Small Versus Large Intestine

E. T. Moran Jr.1

Department of Poultry Science, Auburn University, Auburn, AL 36849-5416

Primary Audience: Microbiologists, Nutritionists, Researchers

SUMMARY
Nutrient absorption occurs from the intestinal surfaces. Strategy in nutrient recovery by the
small intestine is opposite to that of the large intestine. The duodenum, jejunum, and ileum have
a very expansive mucosa by virtue of villi from the wall and microvilli on enterocytes, whereas
the cecal body dominates the large intestine with its mucosa providing minimal surface area
exposure to the lumen. Refluxive motility and villi movement in the small intestine provide ready
convective contact between lumen contents and the unstirred water layer at the villi surfaces while
contractile elements move microvilli to enhance contact after transfer into this surface. Refluxive
motility in the colon gently separates fine from coarse particulates of small intestinal indigesta
using urine moved from the urodeum. Cecal entry is restricted to fluid and fines by narrow orifices
and protruding villi. Microbes in the small intestine are suppressed and reflect a bird’s environment,
whereas rapid motility favors aerobes because of oxygen exchange with the wall. Microbes in the
large intestine are highly concentrated with low oxygen levels that support anaerobes. Plant fiber
that has high proportions of cellulose lends to coarse particulates after feed manufacturing, whereas
fiber high in hemicellulose-pectin combinations disintegrate and may be partially soluble. Fiber
generally acts as a dietary diluent with coarse particulates being more rapidly evacuated than
soluble nonstarch polysaccharides or fines. Soluble nonstarch polysaccharides that increase viscosity
of the small intestine’s luminal contents adversely affect live performance because of impaired
efficiency of convection, reduced rate of exchange with the mucosa, and an expanded microbial
population. Both nonstarch soluble fiber and the fine particulates are readily fermented to volatile
fatty acids in the ceca that contribute to metabolizable energy.

Key words: fiber, large intestine, intestinal microflora, small intestine

2006 J. Appl. Poult. Res. 15:154–160

DESCRIPTION OF PROBLEM to voiding waste. Motility in the large intestine

The small intestine uses a bird’s own en-
zymes to effect digestion and then rapidly cap-
tures resulting nutrients at its expansive surface.
Small intestinal recovery is heavily dependent
on convection driven by motility to minimize
microbial use. Conversely, the large intestine
subsequently uses an extensive symbiotic micro-
bial population to finalize nutrient recovery prior
is largely involved in segregating fiber sensitive
to microbial action while voiding the coarse ma-
terial. Fiber associated with plant feedstuffs can
exert considerable influence on small and large
intestinal functioning by virtue of its solubility
in water and physical characteristics. The struc-
ture of fiber and its relationship with water in
the lumen greatly influence convective effi-
ciency and microbial dynamics throughout the

1
Corresponding author: emoran@acesag.auburn.edu

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 MORAN: INFORMAL NUTRITION SYMPOSIUM
intestine. The following is an overview of rela-
tionship between fiber and microbes in the small
and large intestinal systems.
SMALL INTESTINE
Motility
Mixing of lumen contents concurrent with
maximizing concentration difference at the in-
testine’s surface is dependent on motility. Initia-
tion of digestion by pancreatic enzymes largely
occurs in the duodenum, whereas the jejunum
and ileum are more involved with nutrient recov-
ery upon release. Differences among these parts
are a function rate of digestion and absorptive
capacity. Motility is essential to convection with
exchanges occurring at 2 distinct areas. Fowl
extensively use the circular muscle layer for re-
fluxive peristalsis to maximize villi exposure [1,
2], whereas fibers originating from muscularis
mucosa rotate each villus, further facilitating dif-
ferences at its surface. Pancreatic enzymes gen-
erally reduce nutrients from large complex forms
to small secondary forms that are particularly
soluble.
Enterocytes finalize digestion at the luminal
surface to simple units for absorption and then
transfer nutrients through the basolateral mem-
brane to the circulatory system. Each enterocyte
has an expanded surface by microvilli with an
attached glycocalyx. Goblet cells provide mucin
that engulfs villus microvilli and their glycoca-
lyx to produce the “unstirred water layer” [3].
Essentially mucin has repeating polypeptide
units held together by cystine, then repeating
units have large amounts of threonine and serine
with connecting carbohydrates to create a bottle-
brush structure [4, 5]. The hydrodynamic associ-
ations of mucin with water and interlinking cre-
ate a micronet that limits sugars, peptides, and
fat micelles to the enterocyte surface. Once mac-
romotility by peristasis and villi rotation has
enabled products of pancreatic enzyme digestion
to enter the enterocyte’s unstirred water layer,
then micromotility of underlying microvilli fa-
cilitates contact to finalize digestion and ab-
sorption.
Surface Adaptation
Villi rapidly and continuously respond to
lumen conditions. Maintenance of the surface
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155
and cost of accommodation is extensive. Villi
can increase or decrease their length to alter
surface area and optimize cost of maintenance
for recovery relative to productive advantage.
All changes originate with proliferation of stem
cells in the crypt. Enterocytes and goblet cells
evolve in proportion to need and then mature to
support surface obligations as they ascend [6].
The intestinal surface arising with embryologi-
cal development is focused on macromolecular
transfer while in ovo. Transition of this surface
to one competent at digestion-absorption coin-
cides with depletion of yolk sac reserves and
self sufficiency [7]. Extensive increases in villi
prominence and surface digestive enzymes oc-
cur through the first 7 d after hatch [8, 9]. Access
to feed at this time has a dramatic effect on the
rate of transition, surface cell proportions, and
nature of mucin in the unstirred water layer [10,
11, 12].
Fiber
The structure of fiber is such that a wide array
of physical conditions may be created within
the small intestinal lumen. Primary cell walls
approximate 90% cellulose, hemicellulose, and
pectic acids with 10% of 3 types of structural
proteins. Cellulose is a very large composite
of glucose polymers that provide the bulk of
structural strength. Xylans are small carbohy-
drate polymers that H-bond to cellulose fibers
with pectic acids acting as an intermediary con-
nection to other xylans and fibers. Glucans ap-
pear to be an additional hemicellulose largely
associated with grain endosperm [13, 14]. As
the necessity for wall strength increases so also
does the proportion of cellulose at the expense
of the other polymers. Seed endosperm cells are
not in need of strength as much as cohesiveness;
thus, hemicelluloses dominate with many being
soluble. As the amount and structural complexity
of these soluble nonstarch polysaccharides differ
among the grains (Table 1) so does the viscosity
of their solutions [15].
Insoluble fiber is generally innocuous during
its journey through the small intestine. If feed
formulation leads to an increase in fiber that
decreases the plane of nutrition then enhanced
motility increases luminal throughput while the
villus lengthens. Increases in the levels of solu-
ble fiber have similar effects, but repercussions
 156 JAPR: Symposium

Table 1. Properties of water-soluble nonstarch polysaccharides from several grains high in concentrations1

Total solids

Analysis Wheat Rye Triticale Barley Oats

Proximate
Yield, % 0.59 1.76 0.69 1.16 0.98
Carbohydrate, mg/g 680 719 673 651 641
CP, mg/g 17.7 8.6 19.9 14.0 18.1
Monosaccharide, mg/g
Rhamnose 2.2 — 2.0 2.7 1.6
Arabinose 237.8 254.3 280.8 65.8 80.8
Xylose 204.1 364.1 266.6 74.8 62.9
Mannose 10.8 18.1 17.9 8.7 3.2
Galactose 122.3 28.0 72.4 18.3 48.9
Glucose 42.9 54.8 33.5 480.7 443.3
Characteristics
Weight average MW2 255,000 770,000 569,000 665,000 446,000
Number average MW2 60,500 90,300 66,000 89,700 66,500
Polydispersity2 4.2 8.5 8.6 7.4 6.7
Viscosity3 1.7 5.9 4.0 4.5 3.1
1
Selected data from Girhammar and Nir [15] on 1 variety of each grain.
2
Molecular weight based on weight and number of polymers while polydispersity is their ratio.
3
Intrinsic viscosity expressed in millipascales relative to water.

may be further encountered if increased viscos-
ity of lumen contents also occurs [16, 17]. In-
creasing viscosity of lumen contents not only
decreases laminar flow and convective effi-
ciency of villi for nutrient absorption, but gas
exchange between wall and digesta also lessens.
A lower partial pressure for oxygen with in-
creased concentrations of nutrients can enhance
development of transient microbes, particularly
anaerobes. Chicks experience digesta viscosity
problems when coliforms, streptococci, and
clostridia dominate the flora early in life until the
more favorable competitive lactobacilli become
competitive [18, 19]. Increasing exposure of co-
liforms concurrent with the displacement of em-
bryo enterocytes from the villus early in life
greatly threatens the chick by their translocation
through imperfections of the surface [20]. Con-
veying an extensive and disproportionate mi-
croflora from the environment compounds im-
pairs nutrient recovery and live performance
when viscous nonstarch polysaccarides are pres-
ent. Chicks have been shown to greatly magnify
the adverse effects of viscous, highly methylated
citrus pectin (Table 2) compared with germ-free
conditions while aberrations in villus shape and
expanded anaerobe numbers are apparent [21,
22].
Supplementation of feed with enzymes that
cleave internal bonds to reduce molecular com-
plexity can greatly reduce viscosity of hemicel-
lulose solutions. Given that xylan structure and
its molecular complexity differ markedly among
the grains so does enzyme specificity for endo-
hydrolysis and viscosity relief [23]. Repercus-
sion of digesta viscosity largely involves in-
creased cost of maintaining an expanded mucosa
in response to an expanded microbial population
that competes for nutrients [24, 25]. Viscosity
per se also does not greatly impair nutrient up-
take with exception of fat; in this case a reduced
frequency of wall contact by the large lipid di-
gestion micelles appears to be at fault [26, 27].
Broad-spectrum antibiotics can relieve the
adverse effects of soluble fiber to the extent of
microbial sensitivity. Similarly, xylanase-gluca-
nase supplements are just as effective as their
compatibility with fiber source. Thus, combina-
tions of enzyme and antimicrobial supplements
are usually better than either one alone but do
not provide an additive improvement [28, 29].
Viscous solutions of nonstarch polysaccharides
impair chick performance to the greatest extent
early in life when their microbial population is
evolving and mucosal surface area represents
the limiting facet of growth rate [30]. Broad-

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 MORAN: INFORMAL NUTRITION SYMPOSIUM 157

Table 2. Effect of highly methylated citrus pectin (HMC) in corn-soybean feed with conventional and germ-free
chicks1

Conventional (1–21 d) Germ-free (3–21 d)

Measurement Corn + HMC Corn + HMC

Live performance
Weight gain, g 773a 693b 737a 694b
Feed per bird, g/d 48.8a 50.9b 51.1 49.5
Water per bird, g/d 139a 181b 154 154
Nutrient utilization
DM digestibility, % 71.6a 64.8b 74.4 74.6
Fat digestibility, % 81.3a 65.7b 93.9 94.8
N retention, % 56.9a 51.9b 62.4 63.8
ME in feed, MJ/kg 13.32b 11.81a 13.76b 14.26a
Digesta viscosity, mPa/s
Duodenum jejunum 1.7b 60.8a 2.2b 7.3a
Ileum 2.4 8.9 2.8b 12.8b
Digesta pH
Duodenum + jejunum 5.3 5.2 6.6 6.5
Ileum 6.6a 5.5b 7.9 7.8
Weight with contents
Total small intestine 66b 88a 4.9 5.3
Ceca 7.9 8.9 8.1b 16.2a
Organic acids
Lactic 1.57b 2.13a — —
Total VFA 0.33 0.42 — —
Ileum villi, %
Zigzag 61a 7b 31 43
Tongue-shaped 21b 53a 33 32
Leaf-shaped 36 42 39 43
Ridge-shaped 36a 1b 24 24
1
Selected data from Langhout et al. [21].

spectrum antibiotics measurably decrease main-
tenance needs of the small intestinal mass to the
advantage of production [31, 32].
LARGE INTESTINE
Motility
Coordinating movement of contents in ceca,
colon, and cloaca is central to effective opera-
tion. The ileocolonic valve intermittently opens
to enable entry of indigesta into the colon. Gentle
refluxive peristasis washes indigesta with urine
from the cloaca while pushing resulting fluids
and fines into the ceca [33, 34, 35, 36]. This
size limitation of particulates is due to the small
entrance of the ceca and projection of villi into
the lumen. The villi profile rapidly diminishes
such that blunting ultimately prevails in the cecal
body to give the appearance of a flat mucosa
[37], and volume in the body is far greater than
in the colon and coprodeum. These blunt villi
have goblet cells largely located in the crevices,
whereas enterocytes predominately face the lu-
men [38].
Cecal enterocytes can actively transport nu-
trients immediately after hatch, but subsequent
turnover of the surface leads to cells being pas-
sive in this respect [39, 40]. Early active trans-
port appears to compensate for a concurrent lim-
ited capacity by embryonic enterocytes in the
small intestine. Although the cecal wall can pas-
sively absorb nutrients, resident microbes pres-
ent an infinitely greater surface area for recovery
than the lumen surface. Thus, nutrient absorption
by the cecal wall largely represents an excess of
microbial need or waste from their metabolism.
Effective cecal functioning depends on a popula-
tion of microbes that are continuously provided
indigesta and held in an anaerobic environment.
Strict anaerobes dominate the population [41,
42, 43], and oxygen limitation depends on a
thick mucus layer [38, 44] with gentle motility
to minimize diffusion from the wall.

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 158 JAPR: Symposium

Ceca Restrictions may be fermented to VFA, its contribution to

Use of soluble fiber and fine particulates en-
hances the efficiency of microbial fermentation.
Entry of urine enables substantial recovery of
associated water and minerals, particularly so-
dium [45], while carrying fiber especially suited
for rapid fermentation to volatile fatty acids [46]
and waste nitrogen when dietary crude protein
is lacking [47]. Coarse plant particulates have
generally maintained their integrity because of
high proportions of cellulose, whereas high pro-
portions of hemicellulose and pectins permit dis-
integration. In turn, feedstuffs that generate fine
particulates with grinding and feed manufactur-
ing practices generally favor entry into the ceca
and energy recovery as volatile fatty acids
(VFA) [48, 49]. The caloric value of nonstarch
polysaccharides (and undigested starch) recov-
ered as VFA approximates 1 kcal/g or 25 to 35%
of the original monosaccharide [50]. Although
a large part of the neutral detergent fiber in feed
metabolizable energy varies considerably with
plant source and sugars involved [51, 52, 53, 54].
Ceca are inoculated after the chick’s emer-
gence from the shell with microflora from the
environment, and then representative member-
ship progressively matures as access to fiber
and anaerobic conditions develop. Duration to
competence at fermentation and meaningful for-
mation of VFA to metabolizable energy is vari-
able. Inclusion of readily fermentable substrates,
such as lactose, greatly enhances early VFA pro-
duction by chicks [55]. Broad-spectrum antibiot-
ics appear to delay population maturity [56], but
meaningful amounts of VFA can eventually be
attained to suppress Enterobacteriacae number
[57]. Although antibiotics can enhance perfor-
mance of a young bird when extensive microbial
levels occur in the small intestine, any advantage
in terms of the ceca seems minimal given their
variable influences on the population and rate
of fermentation [58].

CONCLUSIONS AND APPLICATIONS

1. The small intestine has an aggressive motility to facilitate convective exposure of luminal
contents to an expansive surface, whereas motility with the large intestine gently exposes
contents to a minimal surface.
2. Microbes within the small intestine are minimized, and oxygen transfer from the walls discour-
ages anaerobes; conversely, the large intestine makes every effort to maximize numbers that
rely on minimal oxygen.
3. Complex carbohydrates capable of microbial disassembly are largely metabolized within the
ceca to VFA that contribute to feed digestible energy. Soluble fiber that creates viscous conditions
with the small intestinal lumen decreases convective efficiency and oxygenation of contents to
expand anaerobe numbers and confound nutrient recovery.

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