Carbohydrate

Digestion and
Metabolism
Overview of Carbohydrate
Digestion and Metabolism
 Carbohydrates
•Carbohydrates are composed of carbon and water
and have a composition of (CH2O)n.
•The major nutritional role of carbohydrates is to
provide energy and digestible carbohydrates provide
4 kilocalories per gram.

energy
Carbon dioxide
Water GLUCOSE
Chlorophyll

6 CO2 + 6 H20 + energy (sun)C6H12O6 + 6 O2

Simple Sugars -
Disaccharides
 Complex carbohydrates
 Oligosaccharides
 Polysaccharides
 Starch
 Glycogen
 Dietary fiber (Dr. Firkins)
Starch
 Major storage carbohydrate
in higher plants
 Amylose – long straight
glucose chains (α1-4)
 Amylopectin – branched
every 24-30 glc residues (α
1-6)
 Provides 80% of dietary
calories in humans
worldwide
Glycogen
G
 Major storage
G G
G G G
carbohydrate in G G
animals G G
G a 1-6 link
G GG
 Long straight glucose
a 1-4 link G G
chains (α 1-4) G
 Branched every 4-8 glc
G
G
residues (α 1-6)
 More branched than
starch
 Less osmotic pressure
 Easily mobilized
 Digestion
 Pre-stomach – Salivary amylase : a 1-4
endoglycosidase
G
G G
G
G G G G G GG a Limit dextrins
G GG
G amylase
G
G G
G a 1-6 link GGG
GG G
maltotriose
GG
a 1-4 link G
G GG
G maltose G
G
isomaltose
Stomach
 Not much carbohydrate digestion
 Acid and pepsin to unfold proteins
 Ruminants have forestomachs with
extensive
microbial populations to breakdown and
anaerobically ferment feed
Small Intestine
 Pancreatic enzymes
a-amylase
maltotriose maltose
G G G G G G G G + G G

a amylase
amylose

G G G G G G G G
GG GG G G
GG G
amylopectin
a Limit dextrins
Oligosaccharide digestion..cont
a Limit dextrins G
G G G
GG G sucrase
G G

maltase G
G Glucoamylase (maltase)
or G G G
a-dextrinase
G G a-dextrinase
GG G G
G
GG G
G
Small intestine
Portal for transport of virtually
all nutrients

Water and electrolyte balance

Enzymes associated with
intestinal surface membranes
i. Sucrase
ii. a dextrinase
iii.Glucoamylase (maltase)
iv.Lactase
v. peptidases
Carbohydrate absorption

Hexose transporter

apical basolateral
Carbohydrate

Comparative Ruminant vs.

Non-Ruminant Animal
 Digestion and Absorption
Non-ruminant Ruminant
CHO in feed
digestive microbial
enzymes fermentation

Glucose in Volatile fatty acids

small intestine in rumen

Absorption into
blood circulation
Digestion of Carbohydrates
 Monosaccharides
 Do not need hydrolysis before absorption
 Very little (if any) in most feeds
 Di- and poly-saccharides
 Relatively large molecules
 Must be hydrolyzed prior to absorption
 Hydrolyzed to monosaccharides

Only monosaccharides can be absorbed

Non-Ruminant Carbohydrate
Digestion
 Mouth
 Salivary amylase
 Breaks starches down to maltose
 Plays only a small role in breakdown because
of the short time food is in the mouth
 Ruminants do not have this enzyme
 Not all monogastrics secrete it in saliva
 Carbohydrate Digestion
 Pancreas
 Pancreatic amylase
 Hydrolyzes alpha 1-4 linkages
 Produces monosaccharides, disaccharides,
and polysaccharides
 Major importance in hydrolyzing starch and
glycogen to maltose
Amylase
Polysaccharides Disaccharides
 Digestion in Small Intestine
 Digestion mediated by enzymes
synthesized by cells lining the small
intestine (brush border)

Brush Border Enzymes

Disaccharides Monosaccharides

* Exception is β-1,4 bonds in cellulose

 Digestion in Small Intestine
Sucrase
Sucrose Glucose + Fructose
* Ruminants do not have sucrase

Maltase
Maltose Glucose + Glucose

Lactose Lactase Glucose + Galactose

* Poultry do not have lactase
 Digestion of Disaccharides

 Newborns have a
full complement of
brush-border
enzymes

Miller et al. (eds.), 1991

 Digestion in Large Intestine
 Carnivores and omnivores
 Limited anaerobic fermentation
 Bacteria produce small quantities of cellulase
 SOME volatile fatty acids (VFA) produced by
microbial digestion of fibers
 Propionate
 Butyrate
 Acetate
Digestion in Large Intestine
 Post-gastric fermenters (horse and
rabbit)
 Can utilize large quantities of cellulose
 Cecum and colon contain high numbers of
bacteria which produce cellulase
 Cellulase is capable of hydrolyzing the
beta 1,4- linkage
 Overview Monogastric
Carbohydrate Digestion
Location Enzymes Form of Dietary CHO
Mouth Salivary Amylase Starch Maltose Sucrose Lactose

Stomach (amylase from saliva) Dextrin→Maltose

Small Intestine Pancreatic Amylase Maltose

Brush Border Enzymes Glucose Fructose Galactose

+ + +
Glucose Glucose Glucose

Large Intestine None Bacterial Microflora Ferment Cellulose

Carbohydrate Absorption in
Monogastrics
 With exception of newborn animal (first
24 hours), no di-, tri-, or
polysaccharides are absorbed

 Monosaccharides absorbed primarily in

duodenum and jejunum
 Little absorption in stomach and large
intestine
 Small Intestine

Carbohydrates Monosaccharides

Portal Vein Active

Transport

Distributed to
Liver tissue through
circulation
Nutrient Absorption - Carbohydrate
 Active transport for glucose and
galactose
 Sodium-glucose transporter 1 (SGLT1)
 Dependent on Na/K ATPase pump
 Facilitated transport for fructose
 Carbohydrate Digestion
in Ruminants
 Ingested carbohydrates are exposed to
extensive pregastric fermentation
 Most carbohydrates fermented by microbes

 Rumen fermentation is highly efficient

considering the feedstuffs ingested
 Reticulorumen

 Almost all carbohydrate is fermented in

the rumen
 Some ‘bypass’ starch may escape to the small intestine
 No salivary amylase, but have plenty of
pancreatic amylase to digest starch
Microbial Populations
 Cellulolytic bacteria (fiber digesters)
 Produce cellulase - cleaves β1→4 linkages
 Primary substrates are cellulose and hemicellulose
 Prefer pH 6-7
 Produce acetate, propionate, little butyrate, CO2
 Predominate in animals fed roughage diets
 Microbial Populations
 Amylolytic bacteria (starch, sugar digesters)
 Digest starches and sugars
 Prefer pH 5-6
 Produce propionate, butyrate and sometimes lactate
 Predominate in animals fed grain diets
 Rapid change to grain diet causes lactic acidosis
(rapidly decreases pH)
 Streptococcus bovis
 Microbial Metabolism
Sugars

ADP
Catabolism
ATP

NADP+
in rumen:
VFA NADPH
CO2 Growth
CH4 Maintenance
Heat Replication
Bacterial Digestion of
Carbohydrates
Rumen:
 Microbes attach to (colonize) fiber
components and secrete enzymes
 Cellulose, hemicellulose digested by cellulases and
hemicellulases
 Complex polysaccharides are digested to yield
sugars that are fermented to produce VFA
 Starches and simple sugars are more rapidly
fermented to VFA
 Protozoa engulf starch particles prior to
digesting them
Ruminant Carbohydrate Digestion
 Small Intestine
 Secretion of digestive enzymes
 Digestive secretions from pancreas and liver
 Further digestion of carbohydrates
 Absorption of H2O, minerals, amino acids, glucose,
fatty acids

 Cecum and Large Intestine

Bacterial population ferments the unabsorbed
products of digestion
 Absorption of H2O, VFA and formation of feces
 Summary of Carbohydrate in
Monogastrics

 Polysaccharides broken down to

monosaccharides
 Monosaccharides taken up by active transport or
facilitated diffusion and carried to liver
 Glucose is transported to cells requiring energy
 Insulin influences rate of cellular uptake
 Carbohydrates Metabolism in Monogastrics
 Serve as primary source of energy in the cell
 Central to all metabolic processes
Glucose
Cytosol - anaerobic

Hexokinase

Pentose
Phosphate Glucose-6-P Glc-1- phosphate
Shunt
glycolysis
glycogen

Pyruvate
 cytosol Pyruvate

mitochondria
(aerobic) Aceytl CoA
FATTY ACIDS

Krebs Reducing
cycle equivalents
AMINO
ACIDS

Oxidative
Phosphorylation
(ATP)
 Control of enzyme activity

Rate limiting step

Glucose utilization
Stage 1 – postparandial
All tissues utilize glucose

Stage 2 – postabsorptive
KEY – Maintain blood glucose
Glycogenolysis
Glucogneogenesis
Lactate
Pyruvate
Glycerol
AA
Propionate
Spare glucose by metabolizing fat

Stage 3- Early starvation

Gluconeogenesis

Stave 4 – Intermediate starvation

gluconeogenesis
Ketone bodies

Stage 5 – Starvation
Carbohydrate Metabolism/
Utilization- Tissue Specificity
 Muscle – cardiac and skeletal
 Oxidize glucose/produce and store glycogen (fed)

 Breakdown glycogen (fasted state)

 Shift to other fuels in fasting state (fatty acids)

 Adipose and liver

 Glucose  acetyl CoA

 Glucose to glycerol for triglyceride synthesis

 Liver releases glucose for other tissues

 Nervous system
 Always use glucose except during extreme fasts

 Reproductive tract/mammary
 Glucose required by fetus

 Lactose  major milk carbohydrate

 Red blood cells

 No mitochondria

 Oxidize glucose to lactate

 Lactate returned to liver for Gluconeogenesis

 Carbohydrate Digestion Rate
Composition and Digestion of Carbohydrate Fractions
___________________________________________________________
Composition Rumen Digestion (%/h)
_____________________________________________________
Sugars 200-350
Fermentation and Organic Acids 1-2
___________________________________________________________
Starch 10-40
Soluble Available Fiber 40-60
Pectins
B glucans
___________________________________________________________
Insoluble Available Fiber 2-10
Cellulose
Hemicellulose
___________________________________________________________
Unavailable Fiber (lignin) 0
a
 Carbohydrate Metabolism
in Ruminants
 Ingested carbohydrates are exposed to
extensive pregastric fermentation
 Most carbohydrates fermented by microbes

 Rumen fermentation is highly efficient

considering the feedstuffs ingested
 Volatile Fatty Acids
Microbial Fermentation
Carbohydrates VFA’s

Glucose
 Short-chain fatty acids produced by microbes
- Rumen, cecum, colon
 3 basic types:
O O O
CH3 C CH3 CH2 C CH3 CH2 CH2 C
O– O– O–

Acetic acid (2c) Propionic acid (3c) Butyric acid (4c)

 VFA Formation
2 acetate + CO2 + CH4 + heat

1 Glucose 2 propionate + water

1 butyrate + CO2 + CH4

VFAs absorbed passively from rumen to portal blood

Provide 70-80% of ruminant’s energy needs
Rumen Fermentation
 Gases (carbon dioxide and methane)
are primary byproducts of rumen
fermentation
 Usually these gases are eructated or
belched out - if not, bloat occurs
 Bloat results in a severe distension of
the rumen typically on the left side of
the ruminant and can result in death
Uses of VFA
 Acetate
 Energy
 Fatty acid synthesis
 Propionate
 Energy
 Gluconeogenic – glucose synthesis
 Butyrate
 Energy
 Rumen epithelial cells convert to ketone (beta
hydroxybytyrate)

Proportions produced depends on diet

 VFA Production – Molar Ratios
Forage:Grain Acetate Propionate Butyrate

100:0 71.4 16.0 7.9

75:25 68.2 18.1 8.0
50:50 65.3 18.4 10.4
40:60 59.8 25.9 10.2
20:80 53.6 30.6 10.7
Rumen VFA Profiles
Metabolism of VFA
 Overview
 Acetate and butyrate are the major energy
sources (through oxidation)
 Propionate is reserved for gluconeogenesis
 Acetate is the major substrate for
lipogenesis
 Propionate is also lipogenic (though glucose)
Glucose Requirements
 There is less fluctuation in blood
glucose in ruminants and blood glucose
is lower at 40-60 mg/dl
 Reduced fluctuation due to:
 Eat more constantly than monogastrics
 Continuous VFA production
 Continuous digesta flow
 Continuous gluconeogenesis
Overview of Carbohydrates and Ruminants

Diet Protein Carbohydrate Fat

_____________________________________________

Bacterial Fatty Acids

Protein
Starch VFA

Rumen
Propionate Acetate
Butyrate

_____________________________________________
Blood Amino Acids Fatty Acids
Glucose
_____________________________________________
Tissue Protein Lactose Fat
Carbohydrate Digestion and Absorption
Ruminant vs. Monogastrics
Digestive Feature Ruminant Non ruminant
Salivary amylase Zero High – primates
Moderate – pig
Low - carnivores
Pregastric fermentation High+ Zero in MOST cases
Gastric Very low Very low
Pancreatic amylase Moderate High
in SI
Glucose absorption Zero to High
from SI low
Post SI Low Low to High