Nutrition and digestion

Please read chapter 43 in your text from one end to the other
1
Nutrition

-Carbohydrates, proteins, and lipids

-Essential nutrients
amino acids
fatty acids
minerals
vitamins

-The enough but not too much principle

2
Most animals are heterotrophs (or chemoheterotrophs if
you are either pedantic or taking BIO 2022).

They obtain their nutrients mostly from organic

sources: from plants, from other animals, or from
microbes and fungi.

“I am food, I am food. I am the eater of food,..”

The Upanishads (ancient Hindu text)

What do animals get from these resources?

3
Carbohydrates can be
divided into
monosaccharides (the
most important
nutritional hexoses are
glucose, fructose, and
galactose); disaccharides
(sucrose, maltose, and
lactose); and
polysaccharides (starch and glycogen).
Starch is the primary energy storage
molecule in plants. Glycogen is the most
important polysaccharide in animals.
4
Proteins are polymers of
20 amino acids joined by
peptide bonds.

The tertiary and

quaternary structure of
proteins determines their
function.
5
To remember about lipids
-Triacylglycerides are esters of
glycerol and fatty acids.
-They are good sources of energy
and the energy storage of choice in
animals (why? Two reasons?).
-Important components of biological
membranes (Why?)

6
 Long
And saturated FAs
have high melting
points (lards)

Short
and unsaturated FAs
have low melting points
(oils)
18 Carbons

Fatty acids without double bonds are called saturated. Those with double
bonds are called unsaturated. Melting point is lower in unsaturated FAs
and in short- than in long-chained fatty acids. 7
 TO REMEMBER
-The most important dietary monosaccharides (hexoses) are glucose,
fructose, and galactose.
-The most important dietary disaccharides are sucrose (G-F), maltose
(G-G), and galactose (lactose-glucose).
-Starch (plants) and glycogen (animals) are important dietary
polysaccharides.
-Proteins are polymers of 20 amino acids joined by peptide bonds.
-Lipids (fatty acids, phospholipids, cholesterol, triacylglycerides) are
non-polar molecules. Triacylglycerides are made of fatty acids and
glycerol. Many lipids are amphipathic
-Fatty acids without double bonds are called saturated. Those with
double bonds are called unsaturated. Melting point is lower in
unsaturated than in saturated FAs and in short- than in long-chained
fatty acids.

8
If this flash tour through the major classes of nutrients
left you dizzy you must read chapters 3-6 of your
textbook (This is material covered ad-nauseaum in BIO
1010)

9
 Some nutrients are indispensable (essential)
Indispensable means that the animals cannot synthesize them in
sufficient amounts and hence it must obtain them in food. A nutrient can
be indispensable because the animal lacks the metabolic pathway to make
it, or because it has limited ability to make enough of the nutrient.

BROAD PRINCIPLE

There are 8 indispensable amino acids but a few others are

“conditionally indispensable” (arginine is indispensable in cats
and can be indispensable during wound/burn healing). 10
 There are two essential fatty acids.
They are essential because animals
do not have the ability to place a
double bond beyond carbon 9 of the
fatty acid (counting from the
carboxyl end)

Linseed from flax

Desaturases 11
Essential fatty acids are important
because they are
a) precursors of important signaling
molecules (such as prostaglandins)

and

b) because they maintain the fluidity

of biological membranes.
12
 Melting
Fatty Acid Abbreviation point
Stearic 18:0 70° C
Acid

Oleic C18:1 n9 16° C

Acid

Linoleic C18:2 n6 -5° C

Acid

Stories about reindeer legs, lizards, and hibernators…

13
 Svalbard reindeer (Pond et al. (1993)

More double
bonds
Unsaturation Unsaturation
index 0.8 0.8 index

0.7 0.7

0.6 0.6

0.5 0.5

0.4 0.4

0.3 0.3

Fewer double
outer
inner

femur

hoof
tibia
tarsus
bonds Subcutaneous deposits Hind leg bones

14
 Minerals

Inorganic nutrients usually required in

small amounts

Mineral Source Function

Calcium dairy, legumes, some vegetables skeleton, signaling
Phosphorous dairy, meat, some grains skeleton, nucleic acids
Sulfur animal protein component of some *amino acids
Chlorine table salt acid-base balance, gastric juice
Sodium table salt nerve function, many others
Iron meat, some vegetables hemoglobin, enzyme co-factor
Iodine sea food, iodized salt component of thyroid hormones

(many others: Fluorine, Zinc, Copper, Manganese, Cobalt Selenium, Chromium,

Molybdenum…)

*methionine and cysteine 15

 A human’s chemical formula
H 375 X106 O132 X106 C85.7 X106 N 6.43X106 Ca1.5 X10 6 P1.02 X106
S206 X103 Na183 X10 3 K177 X10 3 Cl127 X10 3 Mg40 X10 3 Si38.6 X10 3 Fe2.68 X10 3 Zn2.11X10 3
Cu 76 I14 Mn13 F13Cr7 Se4 Mo3Co1

Element Hydrogen Oxygen Carbon Nitrogen Other

% 61.8 25.4 9.4 1.4 1.0
16
99%
 BROAD PRINCIPLE
Vitamins
Organic molecules required in small amounts. Vitamins
have very diverse functions….

Water soluble Source Function

Vitamin C fruits (citrus), vegetables collagen synthesis, immunity

Niacin nuts, meat, grains component of NAD+ and NADP+

Lipid soluble
Vitamin A Green and orange vegetables visual pigments, antioxidant

Vitamin D Dairy, egg yolk, fish absorption and use of Ca

and P
Excess of water soluble vitamins is excreted in urine (moderate overdoses are
more or less harmless). Excess of lipid soluble vitamins is stored in fat and
therefore overdoses may result in toxic effects.
17
 Functions: antioxidant,
synthesis of collagen,
synthesis of carnitine
(aids fatty acid entry
into mitochondria, Symptoms of avitaminosis:
biosynthesis of Scurvy. In this condition the
norepinephrine,..,etc.). structure of collagen is
defective and people end up
Sources: fresh fruit with spongy gums, bleeding from
(citrus, rose hips), liver. mucous membranes, liver spots
in legs.

18
19
 Phylogenetic Distribution of Vitamin C Synthesis

Cannot synthesize Vit. C

Bats Can synthesize vitamin C

Anthropoid Primates

All (most other mammals)

20
Guinea Pig
 BROAD PRINCIPLE

The effect of many minerals and vitamins is dependent on

intake. There is an optimal intake level. Eating to little or too
much can have negative effects… The “enough but not too
much” principle applies to much of biology.
21
 Which one of these nutrients is
NOT essential for humans

E
A

B
D
Iron

C L-methionine 22
 Things to Remember
-An indispensable (essential) nutrient is a nutrient than an
animal cannot synthesize by itself.
-There are 8 indispensable amino acids.
-There are 2 indispensable fatty acids (linoleic and linolenic)
-Essential fatty acids are important because they are
precursors of signaling molecules and because they increase
the fluidity of membranes.
-Animals require a bunch of minerals (please recall the
function of iron, sodium, and sulfur).
-Vitamins are organic molecules required in small amounts.
-Water soluble vitamins (VC and Niacin) are excreted in urine.
Lipid solubles are stored in fat and over-ingestion can be
toxic.
-The effect of many minerals and vitamins follows the “enough
but not too much” principle.

23
How do guts work

24
 Digestive Physiology

-The concept of assimilation

-A variety of gut designs

-The human gut and its glands

-An overview of nutrient assimilation

-Lactose intolerance

-Glucose transport

25
 How do animals assimilate nutrients

Assimilation = Digestion + Absorption

to digest is to break up a large molecule into

smaller ones.

Assimilation (most often takes place in the gastrointestinal

tract). There are exceptions… Arachnids inject digestive
enzymes into their prey.

26
Guts!

27
The digestive system in many (but not all) animals is a
saculated tube into which many glands empty their
contents. 28
29
30
31
The HCl helps in the hydrolysis of protein.
Pepsinogen is the precursor of pepsin 32
33
 Nominal area ≈3.3 m2

Addition of folds of Kerkring ≈ 10 m2

Addition of villi to
Folds of Kerkring ≈100 m2

Addition of microvilli to
villi ≈2,000 m2

Absorptive surfaces maximize their area by successive levels

34
of folding.
The cells of the intestine
are called enterocytes

Gr. Enteron = gut

Cytos = cell

35
 To Remember
1) The digestive process consists of four steps: ingestion,
digestion, absorption, and elimination (defecation).
2) Nutrient assimilation consists of 2 steps: digestion and
absorption
3) The GI tract is a saculated tube with many glands.
4) The digestive and absorptive surfaces of the GI tract
often increase their surface area by multiple levels of
folding.
5) Intestinal epithelial cells are called enterocytes

36
Why would a person with a weak esophageal
sphincter complain of “heartburn”?

A. The sphincter is inhibiting the passage of food

into the stomach from the esophagus, causing
the esophagus to swell.
B. The sphincter is not allowing the passage of
bile salts from the esophagus to the small
intestine.
C. The sphincter is allowing regurgitation of
stomach acids into the relatively unprotected
esophagus.
D. All of the above answers apply.
37
Why is it that the small intestine has so much more
surface area than other major digestive organs?

A. Its huge surface area allows production of

sufficient hydrochloric acid for digestion.
B. Because the small intestine is involved in
mixing and breaking up food mechanically.
C. The extra surface area allows the small
intestine to secrete enough enzymes for
digestion.
D. Its huge surface area makes highly efficient
absorption of nutrients into the bloodstream
possible.
38
The process of assimilation can be divided into several
steps:

Luminal (extracellular) digestion ---> membrane digestion

Often, enzymatic digestion has two steps. 39

 This aplies to all types of nutrients EXCEPT lipids

The assimilation process

luminal membrane
physical enzymatic enzymatic uptake/transport
processing digestion digestion

BROAD PRINCIPLE
40
Figure 43-6 Slide 5

tra sti in
c t ve
ge n
di atio
c
Lo
Carbohydrates Lipids Proteins

1. Mouth Salivary amylase Lingual lipase

2. Esophagus
Pepsin
3. Stomach Polypeptides

Pancreatic Bile salts Trypsin

4. Small intestine α-amylase and pancreatic Chymotrypsin
lipase Elastase
Carboxypeptidase
Lumen Monosaccharides Disaccharides Monoglycerides
of small (simple sugars) Trisaccharides Fatty acids Short peptides
intestine Amino acids

DIFFUSION
Cell membrane of epithelial cell
FACILITATED Monoglycerides FACILITATED
DIFFUSION AND Fatty acids DIFFUSION AND
COTRANSPORT COTRANSPORT
Epithelium of
Triglycerides
small intestine Amino acids
Monosaccharides
Chylomicrons (protein-
coated globules)
FACILITATED
FACILITATED DIFFUSION AND
DIFFUSION EXOCYTOSIS COTRANSPORT

To bloodstream To lymph vessels, To bloodstream

then bloodstream
41
Figure 43-6 Slide 1

tra sti in
c t ve
ge n
di atio
c
Lo
Carbohydrates Lipids Proteins

1. Mouth Salivary amylase Lingual lipase

42
Figure 43-6 Slide 2

tra sti in
c t ve
ge n
di atio
c
Lo
Carbohydrates Lipids Proteins

1. Mouth Salivary amylase Lingual lipase

2. Esophagus
Pepsin
3. Stomach Polypeptides

43
Figure 43-6 Slide 3

tra sti in
c t ve
ge n
di atio
c
Lo
Carbohydrates Lipids Proteins

1. Mouth Salivary amylase Lingual lipase

2. Esophagus
Pepsin
3. Stomach Polypeptides

Pancreatic Bile salts Trypsin

4. Small intestine α-amylase and pancreatic Chymotrypsin
lipase Elastase
Carboxypeptidase
Lumen Monosaccharides Disaccharides Monoglycerides
of small (simple sugars) Trisaccharides Fatty acids Short peptides
intestine Amino acids

44
Figure 43-6 Slide 4

tra sti in
c t ve
ge n
di atio
c
Lo
Carbohydrates Lipids Proteins

1. Mouth Salivary amylase Lingual lipase

2. Esophagus
Pepsin
3. Stomach Polypeptides

Pancreatic Bile salts Trypsin

4. Small intestine -amylase and pancreatic Chymotrypsin
lipase Elastase
Carboxypeptidase
Lumen Monosaccharides Disaccharides Monoglycerides
of small (simple sugars) Trisaccharides Fatty acids Short peptides
intestine Amino acids

DIFFUSION
Cell membrane of epithelial cell
FACILITATED Monoglycerides FACILITATED
DIFFUSION AND Fatty acids DIFFUSION AND
COTRANSPORT COTRANSPORT
Epithelium of
Triglycerides
small intestine Amino acids
Monosaccharides
Chylomicrons (protein-
coated globules)

45
Figure 43-6 Slide 5

tra sti in
c t ve
ge n
di atio
c
Lo
Carbohydrates Lipids Proteins

1. Mouth Salivary amylase Lingual lipase

2. Esophagus
Pepsin
3. Stomach Polypeptides

Pancreatic Bile salts Trypsin

4. Small intestine α-amylase and pancreatic Chymotrypsin
lipase Elastase
Carboxypeptidase
Lumen Monosaccharides Disaccharides Monoglycerides
of small (simple sugars) Trisaccharides Fatty acids Short peptides
intestine Amino acids

DIFFUSION
Cell membrane of epithelial cell
FACILITATED Monoglycerides FACILITATED
DIFFUSION AND Fatty acids DIFFUSION AND
COTRANSPORT COTRANSPORT
Epithelium of
Triglycerides
small intestine Amino acids
Monosaccharides
Chylomicrons (protein-
coated globules)
FACILITATED
FACILITATED DIFFUSION AND
DIFFUSION EXOCYTOSIS COTRANSPORT

To bloodstream To lymph vessels, To bloodstream

then bloodstream
46
Error!!!!

Di- and Trisaccharides

are NOT taken up by
the intestinal cells. They
are first broken up by
membrane-bound
enzymes into
monosaccharides.

47
Sucrose

Glucose + Fructose

SUCRASE

48
 luminal membrane
physical enzymatic enzymatic uptake/transport
processing digestion digestion
many
enzymes transport
enzymes proteins
secreted by
attached to attached to
salivary
the brush- the brush-
glands
border of border of
(amylase,
intestinal intestinal
lipase)
cells cells
stomach
(lactase,
(pepsin)
sucrase)
pancreas
(amylase,
trypsin,
lipase)

BROAD PRINCIPLE
49
 To Remember

1) For many nutrients, the process of assimilation consists

of four steps: physical processing, luminal digestion,
membrane digestion, and uptake/transport.

2) The glands that secrete the enzymes that act in the

lumen of the gastrointestinal tract are: the salivary gland,
the stomach, and the pancreas.

3) Disaccharides must be hydrolyzed (broken down) into

monosaccharides

50
 The evolution of lactose tolerance in
Homo sapiens (sapiens?)

-We are mammals.

-Mammals are hairy, warm blooded (endothermic

homeotherms), viviparous vertebrates, that feed
their young on milk.

-Milk is a heterogeneous solution (its composition

varies from species to species) that contains proteins,
Lipids, carbohydrates, electrolytes, and vitamins.

51
 the
“invention”
of milk

Mammals are the only vertebrates that secrete “true” milk

(pigeons, emperor penguins, and flamingos secrete “crop milk”) 52
 Lactose

-Lactose is the primary carbohydrate

in mammalian milk (a few marsupials
secrete oligosaccharides of galactose).
-Lactose is a disaccharide made of galactose
and glucose joined by a 1-4 b bond.
-Lactose is very rare in nature except in milk.

Glucose + Galactose

SGLT1 Lactase
53
 Most adult
mammals are lactose
intolerant.

Babies, of course, are not!

The loss of intestinal

lactase activity follows a
fixed ontogenetic program
(it is independent of diet)

(data for rats)

54
 Two exceptions:

1) Many pinnipeds
(sea lions and
seals) lack
intestinal lactase
activity (why?).

Their milk lacks

2) Certain human ethnic groups lactose!
(N. European caucasians, pastoral
groups of north and central
Africa) retain lactase activity as
adults (less than 10% of
humanity).
BROAD PRINCIPLE 55
56
-lactose intolerance is a genetically determined trait
-It has simple Mendelian genetics (tolerance is dominant,
intolerance is recessive).
-Lactose intolerance is the ancestral condition in humans
-Lactose tolerance evolved twice in humans, how come?

in both cases as a result of

“coevolution” with domestic
ungulates (cows, goats, and camels).

57
 Hypotheses to explain the evolution of lactose
tolerance
Pastoralism (nutritional advantage, sensu stricto. Low
latitudes)

Calcium absorption (high latitudes)

– The low availability of sunlight, and hence low
synthesis of Vit. D is the the selective agent
– Milk has high Calcium content
– Calcium and lactose enhance each other’s
absorption (mechanism unclear).

Lactose Tolerance may have its bio-

cultural origins in the practice of relying
on milk to supplement mother’s milk

58
59
In summary:

Lactose intolerance is an example of a genetic polymorphism

in humans. It is an example of relatively recent (less than
10,000 years) evolutionary change in human populations.
Lactose tolerance evolved in response to an association with
ruminants (cows, goats, and sheep). It is also an example of
convergent evolution in humans.

It is one of many examples of clinically significant ethnic

variation in physiological traits. Other examples are cystic
fibrosis, sickle cell anemia, and adult onset (type II)
diabetes.

60
 Evolution matters!

Understanding how evolution works has profound

consequences for human health and well-being

1859

1809-1882 61
 To Remember
1) Milk is a unique mammalian trait.
2) The main carbohydrate in milk is the disaccharide
lactose (glucose-galactose).
3) Lactose is hydrolyzed by the membrane-bound intestinal
enzyme lactase.
4) Most mammal babies have lactase, but most adults do
not.
5) Exceptions are some pinnipeds (never have lactase) and a
small fraction of Homo sapiens individuals (≈ 10%) who
have it as adults.
6) Lactose intolerance is the ancestral and most frequent
trait in humans, but tolerance has evolved in humans
twice in pastoralist societies.

62
A brief review of material from LIFE 1010

BROAD PRINCIPLE 63
 A brief review of material from LIFE 1010

Simple difussion
(does not saturate, it always takes place down a
concentration gradient... it is "downhill")

Facilitated diffusion
(always mediated by transport proteins. These can be
channels (does not saturate) or transporters (saturate),
it always takes place down a concentration gradient... it is
"downhill").

Active transport
(always mediated by transport proteins, can be "uphill"
(against a concentration gradient), it always requires
energy, can be primary or secondary)

BROAD PRINCIPLE 64
How are glucose and galactose transported

65
Which of the following mechanisms of membrane transport
DOES NOT participate in the uptake of glucose by
enterocytes?

A) Active transport
B) Facilitated diffusion
C) Simple diffusion
D) Co-Transport

66
The chemical ouabain, blocks the flow of K+ into
cells and therefore stops the action of the Na+/K+
ATPase pump. Ouabain

A) has no effect on the intestinal transport of

glucose
B) stops the intestinal transport of glucose
C) increases the intestinal transport of glucose

67
 Fermentative Digestion
(not in your book in any detail)
-The concept of fiber (why the anisomeric carbon in
glucose matters)
-Nutritional symbioses
-The fermentation process
-Foregut and hindgut fermenters
-Foregut fermentation: the multichambered
stomach (rumination and merycism)
-Hindgut fermentation and its consequences.

Isomeric: made of the same components in the same

proportions.

bradley@uwyo.edu
68
Cellulose and starch differ in the form of the bond that joins the glucose
residues. Cellulose is the most abundant molecule on earth. 69
 Why you can eat, but cannot assimilate grass: a case
against vegetarianism in the high plains.
cellulose
Vs

b glycosidic bonds
All vertebrates have a-amylase, but no known
vertebrates have cellulases (many mollusks and
insects do have cellulases). Then how do cows
manage to assimilate a large fraction of the
cellulose that they ingest?

Cellulose is (with hemicellulose, lignin, and pectin) what

animal nutritionists call “fiber”. Chemical (plant cell walls)
 glycosidic bonds and nutritional (“refractory”) definitions of “fiber”.
70
Many herbivores maintain “nutritional symbioses” with:
Anaerobic
Bacteria
Ciliated Protozoans
Fungi

These have cellulases!

Symbiosis = living together

71
 the host gets
volatile fatty acids (waste
cellulose products of bacterial
+ metabolism) + bacterial bodies
source of nitrogen

BROAD PRINCIPLE

bacterial bodies are yummy (rich in protein, essentials,

vitamins, ..., etc.) 72
A few cool factoids....

Human Microbiome ≈ 100 trillion cells

Human cells ≈ 10 trillion cells

73
General message: Fermenting herbivores (and
humans!!) participate in a nutritional/digestive
mutually beneficial symbiosis with anaerobic
microorganisms. The herbivore provides the microbes
with a relatively homeostatic environment and with
nutrients. The microbes assimilate materials (such as
cellulose and hemicellulose) that cannot be digested
by the herbivore and produce metabolites (volatile
fatty acids) that can be used by the herbivore. The
herbivore also assimilates a fraction of all microbes
and hence receive the benefits of a more or less
balanced diet (vitamins, essential amino acids and
fatty acids).

74
 To Remember
1) The cell walls of plant cells contain the polysaccharide
cellulose.
2) Vertebrates do not have cellulases and few animals are
efficient at breaking down cellulose.
3) Thus, animals have established a partnership with
symbiotic fermentative microbes (bacteria, protoctists,
and fungi) that can break cellulose.
4) The microbes get a nice environment and plenty of food,
the animal gets the microbes’ energy-rich waste
products (plus some microbe flesh served on the side).

75
There are two types of mammalian
fermenting herbivores.

-Fore-gut (pre-gastric) fermenters Colobus guereza

-Hind-gut (post-gastric) fermenters

The difference between them is in the

placement of the fermentation
vat/chamber in the GIT. The Phascolarctos cinereus
fermentation chamber houses the
fermentative microorganisms. + horses, rabbits,
voles, geese, grouse,
iguanas,..,etc.
BROAD PRINCIPLE 76
 In foregut fermenters the fermentation
chamber is located in an oral position
(before) relative to the small intestine.

Cecum

In hindgut fermenters
the fermentation
chamber is located in an
aboral (after) position
relative to the small
intestine.
This anatomical difference has
profound physiological and ecological
consequences.
Colon
Hindgut=Cecum+Colon 77
Cows are foregut fermenters

78
Horses are hindgut fermenters

79
 Fig. 8.17
What about humans?

total
Gg

Pp
Hp Pt

Hl
Ss
Hs

Are humans herbivores? Large intestine 80

 Foregut Fermenters
Birds ( hoatzin, only one species)
Marsupials (kangaroos and wallabies*)
Edentates (three toad sloth)
Primates (colobus and langur monkeys)
Artyodactils (hippopotamus, camels+, sheep+, goats+, deer+,
cows+)
Cetaceans (baleen whales, why???? Chitin=shrimp shells)

Minke whale
Balaenoptera acutorostrata

81
Baleen whales feed on krill
 cellulose

N-acetyl-b-D-glucosamine

Chitin (chitobiose)

82
83
In foregut fermenters, the products of fermentation
contribute with a very large fraction of an animal’s
energy budget. But there is variation

Grazers Browsers (concentrate feeders)

Wallaby 42% Duiker 18-40%
Wildebeest 67% Mule deer 23-45%
Cattle 63-90%
Sheep 53-80%

84
 Form and function of the multichambered fermentative
stomach

Reticulorumen Abomasum, 1 < pH < 3

Omasum
pH≈ 7 (!!) Acid digestion of bacterial protein
pH≈ 7 (!!)
fermentation absorption of VFAs
and absorption Filter/pump
of VFAs
One example
of a large
esophagous diversity

Ventricular (esophageal)
grove
Shunts materials (milk!)
directly from the
esophagous to the
omasum.
85
Llama (cows are more complicated!)
 Houses microbes, absorbs
Means “book”
products of fermentation
(folds),
particle
retention.

Absorbs
products of
fermentations
, H20, some
particle
retention
Acid digestion, secretes lisozyme
86
 To Remember
1) Vertebrates can be divided into fore- and hind-gut
fermenters depending on where in the gut is the
fermentative chamber that houses microbes.
2) Remember some examples of each type of fermenter.
3) Some whales are fore-gut fermenters and ferment the
chitin in crustaceans.
4) In many foregut-fermenters, the foregut is divided into
a reticulorumen (fermentation and absorption, pH 7), an
omasum (pH 7), and an abomasum (acidic stomach,
digests bacteria).

87
On what you can and cannot eat….

And the LORD spake unto Moses and

to Aaron, saying unto them,What
should we eat?

11:2: Speak unto the children of

Israel, saying, These are the beasts
which ye shall eat among all the
beasts that are on the earth….
11:3 Whatsoever parteth the hoof,
and is clovenfooted, and cheweth the
cud, among the beasts, that shall ye
eat.
88
 Which ones could the ancient Hebrews eat?

D
A

89
11:3 Whatsoever parteth the hoof, and is
clovenfooted, and cheweth the cud, among the
beasts, that shall ye eat

+
Not cloven hoofed Chews the cud

= No good (to eat…) 90

 Nitrogen metabolism in fermenters

Foregut fermenters are

efficient in their use of
protein.

-They can use bacterial protein

with high biological value and
they recycle nitrogen.

Nitrogen recycling

The deamination of amino acids yields ammonia which is transformed

into urea in the liver and kidney. A fraction of all urea is excreted in
urine, a fraction is delivered directly to the rumen, and a fraction is
delivered to the rumen through saliva. Urea is transformed into
ammonia by bacterial ureases and used to synthesize protein by
bacteria. The cow, then, can assimilate these microorganisms. 91
Many mammals (and birds and reptiles) show hindgut
fermentation. The reliance on fermentation as an energy
source varies significantly (why??).

Herbivores Omnivores
Rabbit 30-40% Pig 9-23%
Wombat 30% Human 6-9%
Howler Monkeys 31% Rat 5%
Horses 30%
Beaver 10%
Porcupine 10%

Hindgut fermentation takes place in either the large

intestine or the caecum (or in both).

92
Hindgut fermenters also show nitrogen recycling.

However, because there is no abomasum after the large

intestine, hindgut fermenters can lose the nutrients
produced by bacteria to feces….

93
To solve this problem many hindgut fermenters eat their own feces.
Many hindgut fermenters use
Coprophagy=feces ingestion
(hamsters, koalas, rats, guinea pigs, lemurs…among others)
and
Cecotrophy =ingestion of cecotrophs (fecal pellets produced by the cecum)
(rabbits, ground squirrels, beavers)

The contents of the cecum are enriched

in bacteria, water, and solubles as a
result of a process called “colonic
separation”.

The fibrous undigested large particles

are shunted to the colon. Water, small
particles, and soluble materials are
transported into the cecum. The
contents of the cecum are voided
regularly and consumed.

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 The Bible as a source of hypothesis…
11:6 And the hare because he
cheweth the cud, but divideth not
the (cloven?) hoof; he is unclean
unto you.
cheweth the cud?
Nope. Eateth its poop!
(doeth not cheweth it)

divideth not the (cloven?) hoof? √

95
 Cecotrophs are rich in bacteria and nutrients

Cecotrophs Feces
Protein 18.6% > 7.0%
Phosphorous 1.54% > 0.98
Bacteria 142 > 31
(1010/g)
VFAs 180 > 45
(mMol/Kg)

Fiber 17.8 < 29.6%

Data for rabbits Yummy!

96
 To Remember
1) Fore-gut fermenters can recycle nitrogen. They can use
waste nitrogen (urea) to “feed” bacteria, and then
assimilate the high quality protein in the bacteria.
2) Hind-gut fermenters also have nitrogen recycling, but to
be able to assimilate the nitrogen “recycled” into high
quality protein by their bacteria, they either eat their
own feces (coprophagy) or by producing specialized
nutritious feces from the cecae called cecotrophs
(ceotrophy).

97
 Differences between fore- and hindgut fermenters

• Hindgut
Foregut
• No direct
Direct availability
availability
of microbial
of microbial
protein
protein
andand
nutrients
nutrients
(essential
(essential
FAs,
vitamins)
FAs, vitamins). Reliance on coprophagy and cecotrophy in some
• species.
Detoxification
•• Availability of dietary
Rumination allows glucose
particle reduction
• Limited availability of dietary glucose

98
Many birds are hindgut fermenters
(e.g. grouse, domestic fowl). Among
birds only the hoatzin has foregut
fermentation.

pH, morphological equivalent

Crop ___ _________
Gizzard ___ __________
Proventriculus ___ _________
A bird that takes the 2 Fs of life earnestly!

We are done digesting! 99

 Important message: do not swallow anything
bigger than yourself!

Please start reading chapter 42 100

 Review Questions
1) What are the two most important dietary monosaccharides? How many carbons do they have?

2) What are the hexose components of sucrose, maltose, and lactose?

3) Starch is the primary storage carbohydrate in _________________, whereas glycogen is the prmary storage
carbohydrate in ___________________.

4) In proteins, amino acids are joined by a ___________________ bond that links a __________ with a __________
group.

5) A triacylglycerol is an esther of three fatty acids and _______________

6) Stearic acid is a saturated fatty acid with a hydrocarbon chain of 18 carbons. Palmitoleic acid is an unsaturated fatty
acid (it has one double bond) with a hydrocarbon chain of 16 carbons. Which one of these fatty acids has a higher melting
point?

7) Define what is meant by an indispensable nutrient. Is glucose an indispensable nutrient?

8) Explain why you can ingest enormous doses of vitamin C but a large dose of vitamin A would be toxic.

9) What is the apical membrane of enterocytes called?

10) Describe in a diagram the steps involved in the assimilation of starch and sucrose.

11) How is lactose assimilated?

12) Does a baby iguana express lactase in its intestinal cells? Does a calf? Does a sparrow?

101
13) The pH of the lumen of the intestine is a) acidic, b) alkaline, or c) neutral.

14) What do you think is the pH optimum of the pancreatic enzyme trypsin?

15) What is the chemical difference between cellulose and starch? What are its biological consequences?

16) Horses are pre- or postgastric fermenters? What are the other terms for pre- and post-gastric fermenter?

17) Why are some whales pre-gastric fermenters?

18) What are the functions and pH of the reticulorumen and abomasums in ruminants?

19) Where are bacteria assimilated in fore-gut fermenters?

20) What on earth is a cecotroph?

21) Please fill up the answers in slide 96 of lectures 5 and 6.

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