Carbohydrate Digestion

Digestion and Absorption

Non-ruminant
CHO in feed digestive enzymes

Ruminant
microbial fermentation

Glucose in small intestine

Volatile fatty acids 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

Polysaccharides

Amylase

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 -1,4 bonds in cellulose

Digestion in Small Intestine

Sucrose
Sucrase

Glucose + Fructose

* Ruminants do not have sucrase

Maltose

Maltase

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
Mouth Stomach Small Intestine

Enzymes
Salivary Amylase (amylase from saliva) Pancreatic Amylase Brush Border Enzymes

Form of Dietary CHO

Starch Maltose Sucrose Lactose DextrinMaltose Maltose 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

Liver

Distributed to 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

Summary of Carbohydrate Digestion and Absorption 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

Carbohydrate Digestion in Ruminants

Ingested carbohydrates are exposed to extensive pregastric fermentation

Most carbohydrates fermented by microbes before they can be exposed to typical gastric and small intestinal enzymes

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 Do not have salivary amylase, but have plenty of pancreatic amylase to digest starch

Microbial Populations

Cellulolytic bacteria (fiber digesters)

Produce cellulase - cleaves 14 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
in rumen:

ATP

NADP+ NADPH
Growth Maintenance Replication

VFA CO2 CH4 Heat

Bacterial Digestion of Carbohydrates

Rumen:

Microbes attach to (colonize) fiber components and secrete enzymes

Protozoa engulf starch particles prior to digesting them

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

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

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

Volatile Fatty Acids

Carbohydrates
Microbial Fermentation

VFAs

Glucose

Short-chain fatty acids produced by microbes

- Rumen, cecum, colon

3 basic types:
O CH 3 C O CH3 CH2 C O O
CH3 CH2 CH2 C 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 ruminants 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

Energy Gluconeogenic glucose synthesis Energy Rumen epithelial cells convert to ketone (beta hydroxybytyrate)

Propionate

Butyrate

Proportions produced depends on diet

VFA Production Molar Ratios

Forage:Grain Acetate Propionate Butyrate

100:0

71.4

16.0

7.9

75:25
50:50

68.2
65.3

18.1
18.4

8.0
10.4

40:60
20:80

59.8
53.6

25.9
30.6

10.2
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 Protein Fatty Acids Starch VFA Acetate Butyrate

Rumen

Propionate

_____________________________________________ Blood Fatty Acids Amino Acids Glucose _____________________________________________ Tissue Lactose Fat Protein

Carbohydrate Digestion and Absorption

Digestive Feature
Salivary amylase

Ruminant
Zero

Non ruminant

High primates Moderate pig Low - carnivores

Zero in MOST cases

Pregastric fermentation High+

Gastric Pancreatic amylase in SI

Glucose absorption from SI Post SI

Very low
Moderate Zero to low Low

Very low High

High Low to High