Rumen
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The rumen, also known as a paunch, forms the larger part of the reticulorumen, which is the first chamber in the alimentary canal of ruminant animals. It serves as the primary site for microbial fermentation of ingested feed. The smaller part of the reticulorumen is the reticulum, which is fully continuous with the rumen, but differs from it with regard to the texture of its lining.
Brief anatomy
The rumen is composed of several muscular sacs, the cranial sac, ventral sac, ventral blindsac, and reticulum.
The lining of the rumen wall is covered in small fingerlike projections called papillae, which are flattened, approximately 5 mm in length and 3 mm wide in cattle. The reticulum (derived from the Latin for net [1]) is lined with ridges that form a hexagonal honeycomb pattern. The ridges are approximately 0.1 - 0.2 mm wide and are raised 5 mm above the reticulum wall. The hexagons in the reticulum are approximately 2–5 cm wide in cattle. These features increase the surface area of the reticulorumen wall, facilitating the absorption of volatile fatty acids. Despite the differences in the texture of the lining of the two parts of the reticulorumen, it represents one functional space.
Stratification and mixing of digesta
Digested food (digesta) in the rumen is not uniform, but rather stratified into gas, liquid, and particles of different sizes, densities, and other physical characteristics. Additionally, digesta does not merely enter and exit the rumen without event, but it is subject to extensive mixing, and travels along complicated flow paths. Though they may seem trivial at first, these complicated stratification, mixing, and flow patterns of digesta are a key aspect of digestive activity in the ruminant and thus warrant detailed discussion.
After being swallowed, food travels down the oesophagus and is deposited in the dorsal part of the reticulum. Contractions of the reticulorumen propel and mix the recently ingested feed into the ruminal mat. The mat is a thick mass of digesta, consisting of partially degraded, long, fibrous material. Most material in the mat has been recently ingested, and as such, has considerable fermentable substrate remaining. Microbial fermentation proceeds rapidly in the mat, releasing many gases. Some of these gases are trapped in the mat, causing the mat to be buoyant. As fermentation proceeds, fermentable substrate is exhausted, gas production decreases, and particles lose buoyancy due to loss of entrapped gas. Digesta in the mat hence goes through a phase of increasing buoyancy followed by decreasing buoyancy. Simultaneously, the size of digesta particles–relatively large when ingested–is reduced by microbial fermentation and, later, rumination. Incomplete digestion of plant material here will result in the formation of a type of bezoar called Phythobezoars. At a certain point, particles are dense and small enough that they may “fall” through the rumen mat into the ventral sac below, or they may be swept out of the rumen mat into the reticulum by liquid gushing through the mat during ruminal contractions. Once in the ventral sac, digesta continues to ferment at decreased rates, further losing buoyancy and decreasing in particle size. It is soon swept into the ventral reticulum by ruminal contractions.
In the ventral reticulum, less dense, larger digesta particles may be propelled up into the oesophagus and mouth during contractions of the reticulum. Digesta is chewed in the mouth in a process known as rumination, then expelled back down the oesophagus and deposited in the dorsal sac of the reticulum, to be lodged and mixed into the ruminal mat again. Denser, small particles stay in the ventral reticulum during reticular contraction, and then during the next contraction may be swept out of the reticulorumen with liquid through the reticulo-omasal orifice, which leads to the next chamber in the ruminant animal's alimentary canal, the omasum.
Water and saliva enter through the rumen to form a liquid pool. Liquid will ultimately escape from the reticulorumen from absorption through the wall, or through passing through the reticulo-omosal orifice, as digesta does. However, since liquid cannot be trapped in the mat as digesta can, liquid passes through the rumen much more quickly than digesta does. Liquid often acts as a carrier for very small digesta particles, such that the dynamics of small particles is similar to that of liquid.
The uppermost area of the rumen, the headspace, is filled with gases (such as methane, carbon dioxide, and, to a much lower degree, hydrogen) released from fermentation and anaerobic respiration of food. These gases are regularly expelled from the reticulorumen through the mouth, in a process called eructation.
Digestion
Digestion in the reticulorumen is a complex process. Digestion occurs through fermentation by microbes in the reticulorumen rather than the animal per se. The reticulorumen is one of the few organs present in animals in which digestion of cellulose and other recalcitrant carbohydrates can proceed to any appreciable degree.
The main substrates of digestion in the reticulorumen are non-structural carbohydrates (starch, sugar, and pectin), structural carbohydrates (hemicellulose and cellulose), and nitrogen-containing compounds (proteins, peptides, and amino acids). Both non-structural and structural carbohydrates are hydrolysed to monosaccharides or disaccharides by microbial enzymes. The resulting mono- and disaccharides are transported into the microbes. Once within microbial cell walls, the mono- and disaccharides may be assimilated into microbial biomass or fermented to volatile fatty acids (VFAs) acetate, propionate, butyrate, lactate, valerate and other branched-chain VFAs via glycolysis and other biochemical pathways to yield energy for the microbial cell. Most VFAs are absorbed across the reticulorumen wall, directly into the blood stream, and are used by the ruminant as substrates for energy production and biosynthesis. Some branched chained VFAs are incorporated into the lipid membrane of rumen microbes. Protein is hydrolysed to peptides and amino acids by microbial enzymes, which are subsequently transported across the microbial cell wall for assimilation into cell biomass, primarily. Peptides, amino acids, ammonia, and other sources of nitrogen originally present in the feed can also be used directly by microbes with little to no hydrolysis. Non-amino acid nitrogen is used for synthesis of microbial amino acids. In situations in which nitrogen for microbial growth is in excess, protein and its derivatives can also be fermented to produce energy, yielding ammonia.
Lipids, lignin, minerals, and vitamins play a less prominent role in digestion than carbohydrates and protein, but they are still critical in many ways. Lipids are partly hydrolysed and hydrogenated, and glycerol, if present in the lipid, is fermented. Lipids are otherwise inert in the rumen. Some carbon from carbohydrate or protein may be used for de novo synthesis of microbial lipid. High levels of lipid, particularly unsaturated lipid, in the rumen are thought to poison microbes and suppress fermentation activity. Lignin, a phenolic compound, is recalcitrant to digestion, through it can be solubilized by fungi. Lignin is thought to shield associated nutrients from digestion and hence limits degradation. Minerals are absorbed by microbes and are necessary to their growth. Microbes in turn synthesize many vitamins, such as cyanocobalamin, in great quantities—often great enough to sustain the ruminant even when vitamins are highly deficient in the diet.
Microbes in the reticulorumen
Microbes in the reticulorumen include bacteria, protozoa, fungi, archaea, and viruses. Bacteria, along with protozoa, are the predominant microbes and by mass account for 40-60% of total microbial matter in the rumen. They are categorized into several functional groups, such as fibrolytic, amylolytic, and proteolytic types, which preferentially digest structural carbohydrates, non-structural carbohydrates, and protein, respectively. Protozoa (40-60% of microbial mass) derive their nutrients through phagocytosis of other microbes, and degrade and digest feed carbohydrates, especially starch and sugars, and protein. Although protozoa are not essential for rumen functioning, their presence has pronounced effects. Ruminal fungi make up only 5-10% of microbes and are absent on diets poor in fibre. Despite their low numbers, the fungi still occupy an important niche in the rumen because they hydrolyse some ester linkages between lignin and hemicellulose or cellulose, and help break down digesta particles. Rumen Archaea, approximately 3% of total microbes, are mostly autotrophic methanogens and produce methane through anaerobic respiration. Most of the hydrogen produced by bacteria, protozoa and fungi is used by these methanogens to reduce carbon dioxide to methane. The maintenance of low partial pressure of hydrogen by methanogens is essential for proper functioning of the rumen. Viruses are present in unknown numbers and do not contribute to any fermentation or respiration activity. However, they do lyse microbes, releasing their contents for other microbes to assimilate and ferment in a process called microbial recycling, although recycling through the predatory activities of protozoa is quantitatively more important.
Microbes in the reticulorumen eventually flow out into the omasum and the remainder of the alimentary canal. Under normal fermentation conditions the environment in the reticulorumen is weakly acidic and is populated by microbes that are adapted to a pH between roughly 5.5 and 6.5; since the abomasum is strongly acidic (pH 2 to 4), it acts as a barrier that largely kills reticulorumen flora and fauna as they flow into it. Subsequently, microbial biomass is digested in the small intestine and smaller molecules (mainly amino acids) are absorbed and transported in the portal vein to the liver. The digestion of these microbes in the small intestine is a major source of nutrition, as microbes usually supply some 60 to 90% of the total amount of amino acids absorbed. On starch-poor diets, they also provide the predominant source of glucose absorbed from the small intestinal contents. Under conditions of ruminal acidosis, when the environment of the reticulorumen has become too acidic (usually due to excessive fermentation of starches and sugars into VFA and lactate), microbes that favor a lower pH may start to dominate the ecosystem of the reticulorumen. This gives rise to rumen acidosis and often feed intake of the ruminant will drop.
References
- Cronjé, P.; E.A. Boomker (2000). Ruminant Physiology: Digestion, Metabolism, Growth, and Reproduction. Wallingford, Oxfordshire, UK: CABI Publishing. ISBN 0-85199-463-6.
- Dijkstra, J.; J.M. Forbes; J. France (2005). Quantitative Aspects of Ruminant Digestion and Metabolism, 2nd edition. Wallingford, Oxfordshire, UK: CABI Publishing. p. 736 pages. ISBN 0-85199-814-3.
- Hobson, P.N.; C.S. Stewart (1997). The Rumen Microbial Ecosystem, 2nd edition. New York: Springer. ISBN 0-7514-0366-0.