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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

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

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

Corresponding author:

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

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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

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
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
Selected data from Girhammar and Nir [15] on 1 variety of each grain.
Molecular weight based on weight and number of polymers while polydispersity is their ratio.
Intrinsic viscosity expressed in millipascales relative to water.

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

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

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
Selected data from Langhout et al. [21].

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

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Ceca Restrictions may be fermented to VFA, its contribution to

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


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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