Digestion in Ruminants

Ruminants

2.8 billion domesticated ruminants ungulates Pregastric fermentation 4 compartment stomach

reticulum rumen omasum abomasum

Reticulum

Honeycomb lining Formation of food bolus Regurgitation initiated here Collects hardware (nails, wire)

Rumen

Digestion and fermentation vat Contains anaerobic microbes Papillae lining Absorption of VFA

Omasum

Laminae/manyply lining

muscular folds

Reduces particle size Absorption of water Absorption of VFA

Abomasum

True gastric stomach Proteolytic enzymes Gastric digestion Decreased pH from 6 to 2.5

Denatures proteins Kills bacteria and pathogens Dissolves minerals (e.g., Ca3(PO4)2)

Omasum and Abomasum

Rumen Fermentation

Worlds largest commercial fermentation

100 billion liters in domestic animals 1010 to 1012 cells/mL 200 liters (50 gallons) in cows

Ruminants

Continuous culture fermenters

input and output

Lignocellulosic substrates used 8 x 1015 mouths to feed

Rumen Environment

pH 6.0 7.0 Highly reduced 10 15% dry matter 39C 260 280 mOsm

Rumen Microbes

Bacteria

>200 species with many subspecies

25 species at concentrations >107/mL

1010 to 1012 cells/mL 99.5% obligate anaerobes

Rumen Microbes

Protozoa

Large (20-200 microns) unicellular organisms Prey on bacteria Numbers affected by diet

Rumen Microbes

Fungi

Known only for about 20 years Numbers usually low Digest recalcitrant fiber

Symbiotic Relationship

Microbes provide to the ruminant

Digestion of cellulose and hemicellulose Provision of high quality protein Provision of B vitamins Detoxification of toxic compounds

Microbes to Ruminants

Digestion of cellulose and hemicellulose

Cellulases are all of microbial origin Without microbes, ruminants would not be able to use forage crops such as pasture, hay or silage

Microbes to Ruminants

Provision of high quality protein

50-80% of absorbed N is from microbes

Improved microbial efficiency will provide more microbial protein Can get over 3 kg of microbial protein per day Amino acid pattern is very similar to that required by the ruminant animal

High biological value protein source

Microbes to Ruminants

Provision of B vitamins

Meets the ruminants requirements under most conditions

Niacin may be beneficial in early lactation dairy cows

Microbes to Ruminants

Detoxification of toxic compounds

Example

Mimosine in Leucaena causes problems

poor growth, reproduction and hair loss

Hawaiian ruminants, but not those from Australia, have microbes that degrade mimosine so Leucaena could be fed

Transferred rumen fluid to Australia Inoculated rumen Fed Leucaena

Symbiotic Relationship

Ruminants provide to microbes

Housing Garbage removal Nutrients Neutral environment

Ruminants to Microbes

Housing

Reliable heat

39 2C

Guaranteed for 18 to 96 hours depending on diet and type of animal

Straw-fed water buffalo longest rumen residence time Small selective browsers (mouse deer or duiker) shortest time

Ruminants to Microbes

Garbage removal

Absorption of VFA

Energy to ruminant CO2 and CH4

Eructation

Passage of indigestible residue and microbes to lower GI tract

Ruminants to Microbes

Nutrients

Animal eats Saliva provides urea (N source for bacteria)

Ruminants to Microbes

Neutral environment

pH 6.5 to 7.0 Saliva contains bicarbonate and phosphate buffers

Cows produce up to 46 gallons of saliva daily Added during eating and rumination Cow ruminates 10-12 hours/day

Ruminants to Microbes

Neutral environment

If pH 5.7 rather than 6.5

50% less microbial synthesis Rate of carbohydrate use is decreased More lactate and less acetate is produced Further downward pH spiral

In concentrate selectors (like deer), parotid salivary glands are 0.3% of body weight

Rumination

10 12 hours/day Reduces particle size

only small particles leave reticulorumen

Increases surface area for microbial fermentation Breaks down impervious plant coatings

Bacterial Digestion of Protein

Microbes utilize N, amino acids and peptides for their protein synthesis Microbes convert dietary proteins into their own proteins

some amino acid conversion occurs so dietary amino acids does not equal amino acids leaving the rumen

Bacterial Digestion of Lipid

Microbial lipases act on triglycerides Biohydrogenation

Addition of H across double bond to saturate unsaturated fatty acids

Lipolysis
+ 3H20 + Lipases

Esterified Plant Lipid

Free Fatty Acids

Biohydrogenation
Fatty acid 16:0 18:0 18:2 18:3 (palmitic) (stearic) (linoleic) (linolenic) Weight percent of fatty acids Diet Abomasal digesta 26 6 17 31 29 45 4 6

Sheep fed alfalfa hay

Biohydrogenation

Reduction of double bonds Result: fatty acids that are more saturated with hydrogen
Unsaturated

Saturated

Biohydrogenation
18:2 100 con. 18:2 t11 18:1 18:0

18:2 converted (%)

80 60 40 20 0 0 1 2 3 4 5 6

Time (h)
(adapted from Harfoot et al., 1973)

Biohydrogenation of Linoleic Acid

Linoleic acid

isomerase
cis-9, trans-11 CLA reductase trans-11 18:1 reductase
18:0

Factors that Reduce Microbial Growth

Rapid, dramatic ration changes

Takes 3-4 weeks for microbes to stabilize

Feed restricted amounts of diet Feed lots of unsaturated fat

Bacteria do not use fat for energy Inhibit fiber digestion and microbial growth Different types of fat have different effects

Factors that Reduce Microbial Growth

Feed lots of non-structural carbohydrate to lower rumen pH (rumen acidosis)

Slug feeding Feed barley or wheat To prevent acidosis, must balance lactate users and producers

Bacteria and pH Tolerance

Species Type pH

Ruminococcus flavefaciens fiber 6.15 Fibrobacter succinogenes fiber 6 Megasphaera elsdenii lactate user 4.9 Streptococcus bovis lactate producer 4.55

Factors that Maximize Microbial Growth

Maximum dry matter intake Balanced carbohydrate and protein fractions

Bacteria need both energy and N for amino acid synthesis

Gradual ration changes Maintain rumen pH Keep feed available at all times

Why Worry about Rumen Microbes?

Microbes make ruminants less efficient

Aerobic fermentation
Glucose + O2 ATP + CO2 + H2O

Anaerobic fermentation
acetic acid + propionic acid + butyric acid + CO2 + H2O + CH4 + Heat

Glucose