You are on page 1of 18

7

Digestion and
Absorption
The process of digestion simple sugars—glucose,
breaks down the large, galactose, and fructose.
complex molecules of Protein molecules are
many nutrients into their hydrolyzed to single
simplest, most soluble amino acid units and
forms so that absorption some dipeptides. As
and use by the body can dietary nutrients are
take place (Table 7-1). digested, they are
The two basic types of transported through the
action involved in this digestive tract by a
process are mechanical series of contractions of
digestion and chemical the muscular walls of
(or enzymatic) digestion. the gastrointestinal tract.
Mechanical digestion The process of digestion
involves the physical and absorption begins
mastication (chewing), when food first enters
mixing, and movement the mouth and ends with
of food through the the excretion of waste
gastrointestinal tract. products and undigested
Chemical digestion food particles in the
involves splitting the feces (Figures 7-1 and 7-
chemical bonds of 2).
complex nutrients
through enzymatically
catalyzed hydrolysis. Digestion and
The three major types of absorption begin in
foods that require the mouth, with
digestion are fats, the mastication
carbohydrates, and (chewing) of food
proteins. Before and its mixture
absorption takes place, with saliva.
most of the fat in food is Digestion
hydrolyzed to glycerol, continues
free fatty acids (FFAs), throughout the
and some gastrointestinal
monoglycerides and system and ends
diglycerides. Complex with the excretion
carbohydrates are of waste products
broken down to the and undigested
46 Canine and Feline Nutrition
food particles in mouth. These teeth are
the feces. associated with an
increased capacity to
chew and crush food,
which is indicative of
MOUTH
In all species, the mouth TABLE 7-1 DIGESTIVEENDPRODUCTSOF
functions to bring food CARBOHYDRATE,PROTEIN,
into the body, initiate
ANDFAT
physical mastication,
Nutrient Enzymes
and mix the food with
saliva. Saliva is secreted End products
Carbohydrate Amylase Glucose
in response to the sight
and smell of food. It acts Lactase Galactose
as a lubricant to Sucrase Maltase Fructose
facilitate both chewing
and swallowing and also Protein Dipeptidase Dipeptides
serves to solubilize the
Amino peptidase Single amino
dietary components that
stimulate the taste buds Pepsinogen acids
and impart flavor to Pepsin
food. In addition to its Nucleotidase
function in digestion, Nucleosidase
saliva is also important Trypsin
to the dog (and less so in Chymotrypsin
cats) for evaporative Carboxypeptidase
cooling. Compared to Nuclease
many ruminant and Fat Intestinal lipase Glycerol
herbivorous species that Pancreatic lipase Free fatty acids
thoroughly masticate Monoglycerides,
their food, dogs and cats diglycerides
often swallow large 45
boluses of food with
little or no chewing.
However, there are also
important differences
between dogs and cats.
Although domesticated
dogs and cats have the
same number of incisor
and canine teeth (six
incisors and two canines
on both the top and
bottom jaws), the dog’s
mouth contains more
premolars and molars
than does the cat’s
Digestion and Absorption 47

Figure 7-1 Gastrointestinal system of the dog.
a diet containing a larger
proportion of plant
material. Thus the
dentition of dogs is
suggestive of a more
omnivorous diet than is
the dentition of cats,
which is more typical of
the pattern seen in most
obligate carnivores.1
Although both dogs and
cats are considered to be
“meat-eaters,” the dog
has evolved to consume
a diet that is more
omnivorous in nature
than that of the cat.
48 Canine and Feline Nutrition
Another important flavors). Dogs and cats
role of the mouth is its possess taste systems
importance in taste that are consistent with
perception (gustation). the general pattern seen
Taste refers to the in other carnivorous
sensation that arises species. Early studies
from stimulation of the classified the cat’s taste
taste buds, spherical or system into four general
ovoid clusters of types of units (I, II, IIA,
papillae that are located and IIB).2 A similar and
on the surface of the closely corresponding
tongue. Taste receptor set of units has been
cells are located at the identified in the
tip of each taste bud and domestic dog.3 In
are classified into five general, both dogs and
general receptor types: cats are highly sensitive
sweet (sugars), sour to the tastes of amino
(acids), salty, bitter acids and to various
(alkaloids, peptides), types of organic acids
and umami and nucleotides. All of
(monosodium these are substances
glutamate, disodium found in abundance in
guanylate, “meaty” animal tissues.4 Another

Figure 7-2 Gastrointestinal system of the cat.
Digestion and Absorption 49
similarity between dogs species. In lieu of these
and cats is that neither receptors, cats possess a
species exhibits a strong receptor that is highly
preference for salt sensitive to quinine,
solutions.5 Although tannic acid, and
there is evidence that alkaloids—flavors that
salty flavors may are thought to be
enhance the perceived as bitter.9 In
attractiveness of some contrast, dogs are
foods for dogs and that typically repelled by
cats show a slight most bitter flavors.
preference for salt at There is speculation that
relatively high these differences
concentrations, these between dog and cat
species do not taste systems reflect the
demonstrate the “salt cat’s evolutionary
appetite” that is reported adherence to a more
in other omnivores and strictly carnivorous
herbivorous species. feeding pattern, leading
Dogs and cats also to taste preferences for
have several interesting foods derived from only
differences in taste animal tissue, and a loss
receptors. One of the of the ability to perceive
most dramatic is in their flavors that are more
sensitivity to the taste of typically found in fruits
sweet. Dogs, but not and vegetables. In
cats, are sensitive to and contrast, the dog
show preferences for evolved from a canid
sweet foods.6,7 (This is species that, although
one of the reasons that also a predator, was
theobromine toxicity, as more omnivorous in
a result of chocolate nature. Further, the
ingestion, is a significant process of domestication
risk to pet dogs but not selected for a more
to pet cats.) Recent varied omnivorous diet
studies have shown that in dogs, as scavenging
one of two receptor gradually replaced
genes known to encode cooperative hunting as a
for the sweet-taste primary feeding
receptors in taste buds is behavior.10 The impact
not expressed in cats.8 that taste and other
This defect may be special sense
responsible for the lack perceptions such as
of response to sweet odors and food textures
flavors observed in the have upon food selection
domestic cat and in and preferences in dogs
several other felid and cats are discussed in
50 Canine and Feline Nutrition
detail in Chapter 19 (see movements of the
pp. 191-193). esophagus. It then
immediately constricts
after food has passed to
Dogs show a
prevent reflux of the
preference for
stomach contents back
sweet foods, while
cats do not.
into the lower
Conversely, cats esophagus.
(but not dogs) are
attracted to flavors
that are typically STOMACH
perceived as bitter. The stomach acts as a
These differences reservoir for the body,
may reflect the allowing food to be
cat’s evolutionary ingested as a meal rather
adherence to a than continuously
more strictly throughout the day. The
carnivorous proximal section of the
feeding pattern stomach is capable of
and the dog’s
expansion to allow
historically more
storage of large meals, a
omnivorous diet.
function that is assumed
to be of greater
importance for dogs,
ESOPHAGUS who tend to eat large
meals at a given time,
Food passes from the than for cats, who prefer
mouth to the stomach to eat multiple small
through the esophagus. meals per day (see
The cells of the mucosal Chapter 19, pp. 192-
lining of the esophagus 193). In addition to its
secrete mucus in storage function, the
response to the presence stomach also initiates
of food, which further the chemical digestion
aids in lubricating food of protein (and possibly
as it passes to the of fat, in the dog), mixes
stomach. As the food food with gastric
reaches the end of the secretions, and regulates
esophagus, the cardiac the entry of food into the
sphincter, a ring of small intestine. The
muscle at the junction gastric glands, which are
between the esophagus located in the mucosal
and stomach, relaxes to lining of the corpus
allow food to enter the portion of the stomach,
stomach. This ring secrete mucus,
relaxes in response to hydrochloric acid (HCl),
the peristaltic and the proteolytic
Digestion and Absorption 51
enzyme pepsinogen. In secretion of HCl and
dogs, gastric lipase is mucus by the stomach.
secreted throughout the Neurological stimuli are
stomach, but appears to produced in response to
be much less important the anticipation of
for fat digestion than eating, the sight and
pancreatic lipase.11 For smell of food, and the
this reason, it is believed presence of food in the
that the majority of fat stomach. In addition,
digestion still takes psychological stimuli
place in the small such as fear, stress, and
intestine. Mucous anxiety can affect gastric
secretions protect the secretions and
gastric mucosa and also gastrointestinal
lubricate the ingested functioning in animals.
food. HCl is necessary The hormone gastrin is
to maintain a proper pH released in response to
for the occurrence of the presence of food and
enzymatic action. It distention of the
functions to slightly alter stomach. It is produced
the composition of by mucosal glands in the
ingested fat and protein antrum portion of the
in preparation for further stomach. Gastrin
action by digestive stimulates the secretion
enzymes in the small of HCl and mucus and
intestine. Along with also increases gastric
previously formed motility. Another local
pepsin, HCl also hormone,
converts pepsinogen to enterogastrone, is
the enzyme pepsin. This produced by glands
enzyme initiates located in the duodenal
hydrolysis of protein mucosa. Enterogastrone
molecules to smaller is secreted in response to
polypeptide units in the the presence of fat
stomach. The activity of entering the duodenum
pepsin is highest in the and counteracts the
acidic environment of activity of gastrin by
the stomach and is inhibiting acid
reduced and eventually production and gastric
inactivated when food motility.
leaves the stomach and Peristaltic movements
is exposed to the neutral of the stomach slowly
pH of the small mix the ingested food
intestine.12 with gastric secretions,
Both neurological preparing it for entry
and hormonal stimuli are into the small intestine.
important for the The mucosal cells
52 Canine and Feline Nutrition
located in the antral solids, and very high-fat
portion of the stomach meals may cause a
secrete mucus that has a decrease in
more alkaline pH and is stomachemptying rate.
low in digestive Diets that contain
enzymes. Thorough soluble fiber as a fiber
mixing in this portion source cause a decreased
results in the production rate of stomach
of a semifluid mass of emptying when
food called chyme. compared with diets that
Chyme must pass contain insoluble dietary
through the pyloric fiber (see Chapter 2, p.
sphincter to enter the 14, Table 2-1). In
small intestine for addition, there is
further digestion. Like evidence in cats that dry
the cardiac sphincter, the cat foods leave the
pyloric sphincter is a stomach at a slower rate
ring of muscle that is than canned foods,
usually in a constricted except when very small
state. This ring relaxes meals are consumed.13
in response to the strong The shape of the kibble
peristaltic contractions in dry cat food can also
that originate in the influence the rate of
stomach and travel gastric emptying, with
toward the intestine. triangular-shaped pieces
While open, the leaving the stomach at a
sphincter allows small slower rate than round
amounts of chyme to pieces.14
enter the duodenum. The
pyloric sphincter
controls the rate of
passage of food from the
SMALL
stomach into the small INTESTINE
intestine. The rate of
Before reaching the
gastric emptying is
small intestine, most of
affected by a number of
the digestive processes
factors, including the
that occur in dogs and
osmotic pressure,
cats are mechanical in
particle size, and
nature. The chyme that
viscosity of the chyme,
is delivered through the
as well as the degree of
pyloric sphincter to the
gastric acidity and
duodenum is a semifluid
volume. In general, large
mass made up of food
meals have a slower rate
particles mixed with
of emptying than small
gastric secretions.
meals, liquids leave the
Carbohydrates and fats
stomach faster than
are almost unchanged in
Digestion and Absorption 53
composition, but the duodenum. This mucus
protein in the food has protects the intestinal
been partially mucosa from irritation
hydrolyzed to smaller and erosion by the
polypeptides. Even this gastric acids that are
digestion is not crucial, entering from the
however, because the stomach and further
enzymes of the small lubricates the food mass.
intestine are capable of Nutrient Digestion
completely digesting The pancreas and the
intact dietary protein. glands located in the
Therefore the major task duodenal mucosa secrete
of chemical digestion enzymes into the
and the subsequent intestinal lumen that
absorption of nutrients chemically digest fat,
occur in the small carbohydrate, and
intestine. protein. Enzymes
Further mechanical secreted by the intestinal
digestion also occurs in cells include intestinal
the small intestine lipase, amino peptidase,
through the coordinated dipeptidase,
contractions of its nucleotidase,
muscle layers. These nucleosidase, and
movements thoroughly enterokinase. Intestinal
mix the food mass with lipase converts fat to
intestinal secretions, monoglycerides,
increase the exposure of diglycerides, glycerol,
digested food particles and FFAs. Amino
to the mucosal surface, peptidase breaks the
and slowly propel the peptide bond located at
food mass through the the N-terminal of the
intestinal tract. Constant protein molecule, slowly
sweeping motions of the releasing single amino
intestinal villi that line acids from the protein
the surface of the chain. Dipeptidase
mucosa mix the chyme breaks the peptide bond
that is in contact with of dipeptides to release
the intestinal wall and two single amino acid
increase the efficiency units. Both nucleosidase
of absorption of digested and nucleotidase
particles. After food has hydrolyze
entered the small nucleoproteins to their
intestine, large quantities constituent bases and
of mucus are secreted by pentose sugars. Lastly,
the Brunner’s glands, enterokinase converts
which are located inactive trypsinogen, a
immediately inside the proenzyme secreted by
54 Canine and Feline Nutrition
the pancreas, to its neutralize the acidic
active form (trypsin). chyme and provide the
The final digestion of proper pH for the
carbohydrate takes place digestive enzymes to
at the brush border of function.
the small intestine. The Bile is another
cells of the brush border important component of
secrete the enzymes nutrient digestion in the
maltase, lactase, and small intestine. It is
sucrase, which produced by the liver
respectively convert the and stored in the
disaccharides maltose, gallbladder. Bile’s
lactose, and sucrose to primary function in the
their constituent small intestine is the
monosaccharides; emulsification of dietary
glucose, fructose, and fat and the activation of
galactose. Protease certain lipases. These
enzymes secreted by the two processes result in
pancreas include trypsin, the formation of very
chymotrypsin, small, water-soluble
carboxypeptidase, and globules called micelles.
nuclease. Several of The formation of
these are secreted in an micelles results in an
inactive form and are increased surface area
activated by other for the action of lipase
components in the small and also arranges lipid
intestine after release. In molecules into water-
addition, pancreatic miscible forms that are
lipase and amylase are able to gain access to the
released into the aqueous layer covering
intestinal lumen and the microvilli, ultimately
respectively function to facilitating absorption of
hydrolyze dietary fat and fat into the body.
starch into smaller units. Hormonal control of
Cholesterol esterase digestion in the small
secreted by the pancreas intestine involves
catalyzes the formation several components.
of cholesterol esters. Secretin is produced by
Free cholesterol must be the mucosa of the upper
esterified to fatty acids portion of the duodenum
to facilitate its in response to the entry
absorption into the body. of acidic chyme into the
The pancreas also duodenum. It stimulates
secretes a large volume the release of
of bicarbonate salts into bicarbonate from the
the small intestine. pancreas and controls
These salts function to the rate of bile flow
Digestion and Absorption 55
from the gallbladder. approaches the ileal-
Cholecystokinin is also cecal junction. Species
released from this that are typically found
portion of the intestinal in the dog’s small
mucosa in response to intestine include
the presence of fat in the primarily streptococci,
food mass. This lactobacilli, and
hormone stimulates Bifidobacterium spp. in
contraction of the the duodenum and
gallbladder, resulting in jejunum and several
a release of bile into the species of anaerobic
intestinal lumen. bacteria and Escherichia
Cholecystokinin also coli in the ileum.
stimulates secretion of Compared with dogs,
the pancreatic enzymes. healthy cats have
significantly higher
concentrations of
Most of the microbes present in their
important tasks of small intestine, having
chemical digestion duodenal counts as high
and the as 108 CFUs.16 Also,
subsequent species differences are
absorption of found when comparing
nutrients occur in typical bacteria present
the small intestine. in the small intestines of
cats and dogs. The
primary microbes found
Small Intestinal in the duodenum of cats
Microflora include the aerobe
Pasteurella and the
Although of greater
anaerobes Bacteroides,
significance in the large
Eubacteria, and
intestine, the small
Fusobacteria.
intestines of healthy
Microflora present in
dogs and cats also
the small intestine
contain resident
function to prevent
microbial populations.
colonization of
In dogs, bacterial
pathogenic microbes by
numbers in the
competing for available
duodenum and ileum are
nutrients, maintaining an
relatively low, rarely
appropriate lumen
exceeding 104 colony
environment, and
forming units (CFUs)
producing inhibitory
per milliliter (ml).15
compounds. In both
These numbers increase
dogs and cats, small
to approximately 106
intestinal bacteria have
CFU per ml in the distal
also been found to
portion of the ileum as it
56 Canine and Feline Nutrition
produce short-chain monosaccharides,
fatty acids (SCFAs), glycerol, FFAs, and
which may further monoglycerides and
influence the lumen diglycerides. As these
environment and small units are
intestinal health.17,18 In produced, they are
addition, certain absorbed by the body
nutrients and dietary along with dietary
components may vitamins and minerals.
influence small Absorption involves the
intestinal microbial transfer of digested
populations. For nutrients from the
example, intestinal lumen into the
fructooligosaccharides blood or lymphatic
(FOS), a type of system for delivery to
naturally occurring fiber tissues throughout the
that is fermented by body. Like digestion, the
certain intestinal greatest part of
bacteria, have been absorption takes place in
found to enhance the the small intestine.
growth of beneficial The structure of the
bacteria and reduce the inner wall of the small
numbers of potentially intestine is designed to
pathogenic bacteria in provide a high amount
the small intestine of of surface area for the
dogs and cats.19,20 A absorption of nutrients.
similar fiber, The mucosal folds, villi,
mannanoligosaccharides and microvilli of the
(MOS) can also mucosa produce an
beneficially alter small absorptive inner surface
intestinal bacterial area that is
populations, albeit via a approximately 600 times
different mechanism21 that of the outer serosal
(see Chapter 35, pp. layer of the intestine.
467-470 for a complete Villi are fingerlike
discussion). projections that cover
the convoluted folds of
the mucosa. Each
Nutrient Absorption individual villus
In dogs and cats, the contains a vascular
chemical digestion of network of venous and
food is completed in the arterial capillaries and a
small intestine. lymph vessel called a
Digestible protein, lacteal. These function
carbohydrate, and fat are to transport absorbed
hydrolyzed to amino nutrients to either the
acids, dipeptides, portal or lymphatic
Digestion and Absorption 57
circulations. The surface transport of these
of each villus is covered nutrients into the cells.
with numerous, minute In contrast, active
projections called transport involves the
microvilli. These are transport of nutrient
often collectively molecules across the
referred to as the brush intestinal epithelial
border of the small membrane against a
intestine. The cells concentration gradient.
lining the luminal This transport
surface of the villi are mechanism differs from
highly specialized passive diffusion in that
absorptive cells called more energy is required
enterocytes. These cells to transport materials
have a lifespan of only 2 against a concentration
to 3 days, during which gradient. For example,
they absorb nutrients the most common type
from the lumen of the of active transport
small intestine. Old cells mechanism involves a
are continually sloughed membrane protein
off and excreted in the carrier coupled with the
feces, giving these cells active transport of
one of the highest sodium (the sodium
turnover rates of any pump).
tissue in the body. Although some
Nutrient absorption is passive diffusion is
accomplished in the believed to occur, most
small intestine through simple carbohydrates are
several processes. Some absorbed by the body
small molecules are through an active
absorbed by passive process that is linked to
diffusion according to sodium transport and
the osmotic gradient; for uses a specific carrier
example, electrolytes protein. Single amino
and water molecules acids and some
both flow across the dipeptides and
mucosa in response to tripeptides are also
osmotic pressure. absorbed in this manner.
Facilitated diffusion Small peptides that are
involves the transport of absorbed into the cell
large molecules across are immediately
the cellular membrane in hydrolyzed to single
concurrence with the amino acid units before
pressure gradient. being released into the
Carrier proteins located portal circulation.
in the membranes of the Sugars and amino acids
enterocytes facilitate are absorbed into the
58 Canine and Feline Nutrition
villus capillaries and The liver functions to
from there enter the further process the
portal vein, which absorbed
transports these nutrients monosaccharides and
to the liver. Absorption amino acids that arrive
of fat involves the through the portal
interaction of the circulation. Some
fatcontaining micelles monosaccharides are
with the aqueous layer converted to the storage
surrounding the brush carbohydrate form
border. Micelles contain glycogen, and a certain
bile acids, quantity of glucose is
monoglycerides, secreted directly into the
diglycerides, and long- circulation. Some amino
chain fatty acids. acids are released into
Because they are water- the bloodstream, where
miscible, the micelles they circulate to tissues
are able to travel to the for absorption into cells.
brush border, where they Excess amino acids are
are disrupted and their either converted to other
component fat particles nonessential amino acids
are absorbed into the or metabolized by the
cell. The bile remains in liver for energy.
the lumen and Most minerals are
eventually moves down absorbed by the body in
the intestine to be an ionized form. The
reabsorbed and water-soluble vitamins
circulated back to the are transported by
liver. Within the passive diffusion, but
enterocyte, most of the some may be absorbed
fatty acids and glycerol by an active process
are resynthesized into when the diet contains
triglycerides, combined low levels. Vitamin B12
with cholesterol, is unique in its
phospholipid, and requirement for an
protein, and released intrinsic factor for
into the central lacteal as proper absorption (see
either chylomicrons or Chapter 5, p. 34). Fat-
very–low-density soluble vitamins are
lipoprotein (VLDL) made soluble by
transport particles. The combination with bile
central lacteal drains salts and are then
into the major lymph absorbed by passive
vessels, and the particles diffusion through the
eventually enter the lipid phase of the
blood circulation near mucosal cell membrane.
the heart. In general, when there is
Digestion and Absorption 59
normal fat absorption, the length of their large
there is normal fat- intestine is relatively
soluble vitamin short. Relative to body
absorption. size, the dog’s cecum is
not as large as the pig’s,
but it is somewhat larger
LARGE than the cat’s. This
observation is consistent
INTESTINE with the fact that the dog
has adapted to
(COLON) consuming a diet that is
The contents of the more omnivorous in
small intestine enter the nature than that of the
large intestine through cat. The extent to which
the ileocecal valve. The bacterial digestion of
cecum is an intestinal dietary fiber in the
pocket located next to cecum and colon
the junction of the colon contributes to energy
and the small intestine. balance in these species
This portion of the is small compared to the
intestine varies in size contribution for
and functional capacity nonruminant herbivore
among species of species. However,
mammals. The cecum of SCFAs produced by
nonruminant herbivores bacterial fermentation of
such as the horse and fiber are an important
rabbit is relatively large energy source for
and has a highly colonocytes and
enhanced digestive contribute to intestinal
capacity. Likewise, both health.22,23 In addition,
the cecum and large adherent bacteria in the
intestine of the large intestine influence
omnivorous pig are function of the intestinal
enlarged when mucosa and help to
compared with those of regulate the enteric
the carnivorous species. immune system22 (see
Microbial digestion of Section 5, pp. 465-467
dietary fiber in the for a complete
cecum and colon of discussion of the role of
nonruminant herbivores colonic bacteria in
contributes significantly intestinal health).
to the nutrient intake and A chief function of
balance of these the large intestine in
animals. In comparison, dogs and cats is the
carnivorous species such absorption of water and
as the cat and mink have certain electrolytes.
a vestigial cecum, and Unlike the small
60 Canine and Feline Nutrition
intestine, the large References
intestine has no villi and digestion of these
therefore has a lower materials produces
capacity for absorption. various gases, SCFAs,
Although it is able to and other byproducts.
efficiently absorb water When protein reaches
and electrolytes, it has the large intestine in an
no mechanisms for undigested state,
active transport. Along bacterial degradation
with a large volume of results in the production
water, sodium is of the amines indole and
absorbed into the body skatole. In addition,
from the large intestine. hydrogen sulfide gas is
As mentioned produced from the
previously, the bacterial sulfur-containing amino
colonies of the colon are acids of undigested or
capable of digesting poorly digested protein.
some of the indigestible Hydrogen sulfide gas,
fiber and other nutrients indole, and skatole
in the diet that have impart strong odors to
escaped digestion in the fecal matter and
small intestine. The intestinal gas. Certain
products of this bacterial types of carbohydrates
digestion contribute to found in legumes such
the characteristic smell as soybeans are resistant
and color of canine and to digestion by the
feline feces. Undigested endogenous enzymes of
food residues, sloughed the small intestine.
cells, bacteria, and These carbohydrates
unabsorbed endogenous reach the colon and are
secretions make up the metabolized by bacteria,
fecal matter that with a resultant
eventually reaches the production of intestinal
rectum and is excreted gas (flatulence).
from the body. Hydrogen, carbon
Fecal characteristics dioxide, and methane
in dogs and cats can be gases are produced from
significantly affected by the bacterial digestion of
the quantity and type of carbohydrates. A similar
indigestible matter that effect occurs with
is present in the animal’s certain types of fiber.
diet. Bacterial While nonfermentable
fibers resist digestion in
the small intestine and
fermentation in the large
intestine, fermentable
fibers are used as an
Digestion and Absorption 61
energy source by the colons of individual
intestinal bacteria, animals.
resulting in the
production of gases and
SCFAs. The degree to In contrast to the
which flatulence and small intestine, a
strong fecal odors occur primary function of
in dogs and cats that are the large intestine
fed poorly digestible (colon) in dogs and
materials varies with the cats is the
amounts and types of absorption of
materials fed and the water and certain
intestinal flora present in electrolytes,
especially sodium.

1. Morris JG, Rogers QR: Comparative dog and cat nutrition. In


Burger IH, Rivers JPW, editors: Nutrition of the dog and cat,
Cambridge, England, 1989, Cambridge University Press.
2. Boudreau JC, Alev N: Classification chemoresponsive tongue
units of the cat geniculated ganglion, Brain Res 54:157–175,
1973.
3. Boudreau J, White T: Flavor chemistry of carnivore taste systems.
In Bullard RW, editor: Flavor chemistry of animal foods,
American Chemical Society symposium series 67, Washington
DC, 1978, ACS, pp 102–108.
4. Kumazawa T, Nakamura M, Kurihara K: Canine taste nerve
response to umami substances, Physiol Behav 49:875–881, 1991.
5. Yu S, Rogers QR, Morris JG: Absence of salt (NaCl) preference
of appetite in sodium-replete or depleted kittens, Appetite 29:1–
10, 1997.
6. Ferrell F: Preference for sugars and nonnutritive sweeteners in
young Beagles, Neurosci Biobehav Rev 8:199–203, 1984.

7. Houpt KA, Coren B, Hintz HF, Hilderbrant JE: Effect of sex and reproductive status on sucrose preference, food in
body weight of dogs, J Am Vet Med Assoc 174:1083–1085, 1979.
8. Li X, Li W, Wang H, and others: Cats lack a sweet taste receptor, J Nutr 136:1932S–1934S, 2006.
9. Bradshaw JWS: The evolutionary basis for the feeding behavior of domestic dogs (Canis familiaris) and cats (Felis
Nutr 136:1927S–1931S, 2006.
10. Macdonald DW, Carr GM: Variation in dog society: between resource dispersion and social flux. In Serpell J, edito
domestic dog: its evolution, behaviour and interactions with people, Cambridge, England, 1995, Cambridge Univer
pp 199–216.
11. Carriere F, Laugier R, Barrowman JA, and others: Gastric and pancreatic lipase levels during a test meal in dogs, Sc
Gastroenterol 28:443–454, 1993.
12. Smeets-Peeters M, Watson T, Minekus M, Havenaar R: A review of the physiology of the canine digestive tract rela
development of in vitro systems, Nutr Res Rev 11:45–69, 1998.
13. Goggin JM, Hoskinson JJ, Butine MD, and others: Scintigraphic assessment of gastric emptying of canned and dry
healthy cats, Am J Vet Res 59:388–392, 1998.
14. Armbrust LJ, Hoskinson JJ, Lora-Michiels NM, Milliken GA: Gastric emptying in cats using foods varying in fiber
and kibble shapes, Vet Radiol Ultrasound 44:339–343, 2003.
62 Canine and Feline Nutrition
15. Kearns RJ, Hayek MG, Sunvold GD: Microbial changes in aged dogs. In Reinhart GA, Carey DP, editors: Recent a
canine and feline nutritional research: Iams nutrition symposium proceedings, vol 2, Wilmington, Ohio, 1998, Ora
Press, pp 337–351.
16. Johnston K, Lamport A, Batt RM: An unexpected bacterial flora in the proximal small intestine of normal cats, Vet
132:362– 363, 1993.
17. Brosley BP, Hill RC, Scott KC: Gastrointestinal volatile fatty acid concentrations and pH in cats, Am J Vet Res 61:3
2000.
18. Strickling JA, Harmon DL, Dawson KA, Gross KL: Evaluation of oligosaccharide addition to dog diets: influences
digestion and microbial populations, Anim Feed Sci Technol 86:205–219, 2000.
19. Sparkes AH, Papasouliotis K, Sunvold G, and others: Bacterial flora in the duodenum of healthy cats and effect of d
supplementation with fructo-oligosaccharides, Am J Vet Res 59:431–435, 1998.
20. Willard MD: Effects of dietary supplementation of fructo-oligosaccharides on small intestinal bacterial overgrowth
Am J Vet Res 55:654–659, 1992.
21. Ofek I, Beachey EH: Mannose binding and epithelial cell adherence of Escherichia coli, Infect Immunol 22:247–25
22. Buddington RK, Buddington KK, Sunvold GD: The use of fermentable fibers to manage the gastrointestinal tract. I
GA, Carey DP, editors: Recent advances in canine and feline nutritional research: Iams nutrition symposium proce
3, Wilmington, Ohio, 2000, Orange Frazer Press, pp 169–179.
23. Drackley JK, Beaulieu AD, Sunvold GD: Energetic substrates for intestinal cells. In Reinhart GA, Carey DP, editor
advances in canine and feline nutritional research, Iams nutrition symposium proceedings, vol 2 Wilmington, Ohio
Orange Frazer Press, pp 463–472.

You might also like