Ruminants are herbivorous grazing or browsing artiodactyls belonging to the suborderRuminantia that are able to acquire nutrients from plant-based food by fermenting it in a specialized stomach prior to digestion, principally through microbial actions. The process, which takes place in the front part of the digestive system and therefore is called foregut fermentation, typically requires the fermented ingesta (known as cud) to be regurgitated and chewed again. The process of rechewing the cud to further break down plant matter and stimulate digestion is called rumination.[2][3] The word "ruminant" comes from the Latin ruminare, which means "to chew over again".
The first fossil ruminants appeared in the Early Eocene and were small, likely omnivorous, forest-dwellers.[8] Artiodactyls with cranial appendages first occur in the early Miocene.[8]
Phylogeny
Ruminantia is a crown group of ruminants within the orderArtiodactyla, cladistically defined by Spaulding et al. as "the least inclusive clade that includes Bos taurus (cow) and Tragulus napu (mouse deer)". Ruminantiamorpha is a higher-level clade of artiodactyls, cladistically defined by Spaulding et al. as "Ruminantia plus all extinct taxa more closely related to extant members of Ruminantia than to any other living species."[9] This is a stem-based definition for Ruminantiamorpha, and is more inclusive than the crown group Ruminantia. As a crown group, Ruminantia only includes the last common ancestor of all extant (living) ruminants and their descendants (living or extinct), whereas Ruminantiamorpha, as a stem group, also includes more basal extinct ruminant ancestors that are more closely related to living ruminants than to other members of Artiodactyla. When considering only living taxa (neontology), this makes Ruminantiamorpha and Ruminantia synonymous, and only Ruminantia is used. Thus, Ruminantiamorpha is only used in the context of paleontology. Accordingly, Spaulding grouped some genera of the extinct family Anthracotheriidae within Ruminantiamorpha (but not in Ruminantia), but placed others within Ruminantiamorpha's sister clade, Cetancodontamorpha.[9]
Hofmann and Stewart divided ruminants into three major categories based on their feed type and feeding habits: concentrate selectors, intermediate types, and grass/roughage eaters, with the assumption that feeding habits in ruminants cause morphological differences in their digestive systems, including salivary glands, rumen size, and rumen papillae.[18][19] However, Woodall found that there is little correlation between the fiber content of a ruminant's diet and morphological characteristics, meaning that the categorical divisions of ruminants by Hofmann and Stewart warrant further research.[20]
Also, some mammals are pseudoruminants, which have a three-compartment stomach instead of four like ruminants. The Hippopotamidae (comprising hippopotamuses) are well-known examples. Pseudoruminants, like traditional ruminants, are foregut fermentors and most ruminate or chew cud. However, their anatomy and method of digestion differs significantly from that of a four-chambered ruminant.[5]
Monogastric herbivores, such as rhinoceroses, horses, guinea pigs, and rabbits, are not ruminants, as they have a simple single-chambered stomach. Being hindgut fermenters, these animals ferment cellulose in an enlarged cecum. In smaller hindgut fermenters of the orderLagomorpha (rabbits, hares, and pikas), and Caviomorph rodents (Guinea pigs, capybaras, etc.), material from the cecum is formed into cecotropes, passed through the large intestine, expelled and subsequently reingested to absorb nutrients in the cecotropes.
Stylised illustration of a ruminant digestive systemDifferent forms of the stomach in mammals. A, dog; B, Mus decumanus; C, Mus musculus; D, weasel; E, scheme of the ruminant stomach, the arrow with the dotted line showing the course taken by the food; F, human stomach. a, minor curvature; b, major curvature; c, cardiac end G, camel; H, Echidna aculeata. Cma, major curvature; Cmi, minor curvature. I, Bradypus tridactylus Du, duodenum; MB, coecal diverticulum; **, outgrowths of duodenum; †, reticulum; ††, rumen. A (in E and G), abomasum; Ca, cardiac division; O, psalterium; Oe, oesophagus; P, pylorus; R (to the right in E and to the left in G), rumen; R (to the left in E and to the right in G), reticulum; Sc, cardiac division; Sp, pyloric division; WZ, water-cells. (from Wiedersheim's Comparative Anatomy)Food digestion in the simple stomach of nonruminant animals versus ruminants
The primary difference between ruminants and nonruminants is that ruminants' stomachs have four compartments:
The first two chambers are the rumen and the reticulum. These two compartments make up the fermentation vat and are the major site of microbial activity. Fermentation is crucial to digestion because it breaks down complex carbohydrates, such as cellulose, and enables the animal to use them. Microbes function best in a warm, moist, anaerobic environment with a temperature range of 37.7 to 42.2°C (99.9 to 108.0°F) and a pH between 6.0 and 6.4. Without the help of microbes, ruminants would not be able to use nutrients from forages.[22] The food is mixed with saliva and separates into layers of solid and liquid material.[23] Solids clump together to form the cud or bolus.
The cud is then regurgitated and chewed to completely mix it with saliva and to break down the particle size. Smaller particle size allows for increased nutrient absorption. Fiber, especially cellulose and hemicellulose, is primarily broken down in these chambers by microbes (mostly bacteria, as well as some protozoa, fungi, and yeast) into the three volatile fatty acids (VFAs): acetic acid, propionic acid, and butyric acid. Protein and nonstructural carbohydrate (pectin, sugars, and starches) are also fermented. Saliva is very important because it provides liquid for the microbial population, recirculates nitrogen and minerals, and acts as a buffer for the rumen pH.[22] The type of feed the animal consumes affects the amount of saliva that is produced.
Though the rumen and reticulum have different names, they have very similar tissue layers and textures, making it difficult to visually separate them. They also perform similar tasks. Together, these chambers are called the reticulorumen. The degraded digesta, which is now in the lower liquid part of the reticulorumen, then passes into the next chamber, the omasum. This chamber controls what is able to pass into the abomasum. It keeps the particle size as small as possible in order to pass into the abomasum. The omasum also absorbs volatile fatty acids and ammonia.[22]
After this, the digesta is moved to the true stomach, the abomasum. This is the gastric compartment of the ruminant stomach. The abomasum is the direct equivalent of the monogastric stomach, and digesta is digested here in much the same way. This compartment releases acids and enzymes that further digest the material passing through. This is also where the ruminant digests the microbes produced in the rumen.[22] Digesta is finally moved into the small intestine, where the digestion and absorption of nutrients occurs. The small intestine is the main site of nutrient absorption. The surface area of the digesta is greatly increased here because of the villi that are in the small intestine. This increased surface area allows for greater nutrient absorption. Microbes produced in the reticulorumen are also digested in the small intestine. After the small intestine is the large intestine. The major roles here are breaking down mainly fiber by fermentation with microbes, absorption of water (ions and minerals) and other fermented products, and also expelling waste.[24] Fermentation continues in the large intestine in the same way as in the reticulorumen.
Only small amounts of glucose are absorbed from dietary carbohydrates. Most dietary carbohydrates are fermented into VFAs in the rumen. The glucose needed as energy for the brain and for lactose and milk fat in milk production, as well as other uses, comes from nonsugar sources, such as the VFA propionate, glycerol, lactate, and protein. The VFA propionate is used for around 70% of the glucose and glycogen produced and protein for another 20% (50% under starvation conditions).[25][26]
Abundance, distribution, and domestication
Wild ruminants number at least 75 million[27] and are native to all continents except Antarctica and Australia.[4] Nearly 90% of all species are found in Eurasia and Africa.[27] Species inhabit a wide range of climates (from tropic to arctic) and habitats (from open plains to forests).[27]
The population of domestic ruminants is greater than 3.5 billion, with cattle, sheep, and goats accounting for about 95% of the total population. Goats were domesticated in the Near Eastcirca 8000 BC. Most other species were domesticated by 2500 BC., either in the Near East or southern Asia.[27]
Ruminant physiology
Ruminating animals have various physiological features that enable them to survive in nature. One feature of ruminants is their continuously growing teeth. During grazing, the silica content in forage causes abrasion of the teeth. This is compensated for by continuous tooth growth throughout the ruminant's life, as opposed to humans or other nonruminants, whose teeth stop growing after a particular age. Most ruminants do not have upper incisors; instead, they have a thick dental pad to thoroughly chew plant-based food.[28] Another feature of ruminants is the large ruminal storage capacity that gives them the ability to consume feed rapidly and complete the chewing process later. This is known as rumination, which consists of the regurgitation of feed, rechewing, resalivation, and reswallowing. Rumination reduces particle size, which enhances microbial function and allows the digesta to pass more easily through the digestive tract.[22]
Vertebrates lack the ability to hydrolyse the beta [1–4] glycosidic bond of plant cellulose due to the lack of the enzyme cellulase. Thus, ruminants completely depend on the microbial flora, present in the rumen or hindgut, to digest cellulose. Digestion of food in the rumen is primarily carried out by the rumen microflora, which contains dense populations of several species of bacteria, protozoa, sometimes yeasts and other fungi – 1 ml of rumen is estimated to contain 10–50 billion bacteria and 1 million protozoa, as well as several yeasts and fungi.[30]
Since the environment inside a rumen is anaerobic, most of these microbial species are obligate or facultative anaerobes that can decompose complex plant material, such as cellulose, hemicellulose, starch, and proteins. The hydrolysis of cellulose results in sugars, which are further fermented to acetate, lactate, propionate, butyrate, carbon dioxide, and methane.
As bacteria conduct fermentation in the rumen, they consume about 10% of the carbon, 60% of the phosphorus, and 80% of the nitrogen that the ruminant ingests.[31] To reclaim these nutrients, the ruminant then digests the bacteria in the abomasum. The enzyme lysozyme has adapted to facilitate digestion of bacteria in the ruminant abomasum.[32]Pancreatic ribonuclease also degrades bacterial RNA in the ruminant small intestine as a source of nitrogen.[33]
During grazing, ruminants produce large amounts of saliva – estimates range from 100 to 150 litres of saliva per day for a cow.[34] The role of saliva is to provide ample fluid for rumen fermentation and to act as a buffering agent.[35] Rumen fermentation produces large amounts of organic acids, thus maintaining the appropriate pH of rumen fluids is a critical factor in rumen fermentation. After digesta passes through the rumen, the omasum absorbs excess fluid so that digestive enzymes and acid in the abomasum are not diluted.[1]
Tannin toxicity in ruminant animals
Tannins are phenolic compounds that are commonly found in plants. Found in the leaf, bud, seed, root, and stem tissues, tannins are widely distributed in many different species of plants. Tannins are separated into two classes: hydrolysable tannins and condensed tannins. Depending on their concentration and nature, either class can have adverse or beneficial effects. Tannins can be beneficial, having been shown to increase milk production, wool growth, ovulation rate, and lambing percentage, as well as reducing bloat risk and reducing internal parasite burdens.[36]
Tannins can be toxic to ruminants, in that they precipitate proteins, making them unavailable for digestion, and they inhibit the absorption of nutrients by reducing the populations of proteolytic rumen bacteria.[36][37] Very high levels of tannin intake can produce toxicity that can even cause death.[38] Animals that normally consume tannin-rich plants can develop defensive mechanisms against tannins, such as the strategic deployment of lipids and extracellularpolysaccharides that have a high affinity to binding to tannins.[36] Some ruminants (goats, deer, elk, moose) are able to consume food high in tannins (leaves, twigs, bark) due to the presence in their saliva of tannin-binding proteins.[39]
The verb 'to ruminate' has been extended metaphorically to mean to ponder thoughtfully or to meditate on some topic. Similarly, ideas may be 'chewed on' or 'digested'. 'Chew the (one's) cud' is to reflect or meditate. In psychology, "rumination" refers to a pattern of thinking, and is unrelated to digestive physiology.
Methane is produced by a type of archaea, called methanogens, as described above within the rumen, and this methane is released to the atmosphere. The rumen is the major site of methane production in ruminants.[41] Methane is a strong greenhouse gas with a global warming potential of 86 compared to CO2 over a 20-year period.[42][43]
As a by-product of consuming cellulose, cattle belch out methane, there-by returning that carbon sequestered by plants back into the atmosphere. After about 10 to 12 years, that methane is broken down and converted back to CO2. Once converted to CO2, plants can again perform photosynthesis and fix that carbon back into cellulose. From here, cattle can eat the plants and the cycle begins once again. In essence, the methane belched from cattle is not adding new carbon to the atmosphere. Rather it is part of the natural cycling of carbon through the biogenic carbon cycle.[44]
In 2010, enteric fermentation accounted for 43% of the total greenhouse gas emissions from all agricultural activity in the world,[45] 26% of the total greenhouse gas emissions from agricultural activity in the U.S., and 22% of the total U.S. methane emissions.[46] The meat from domestically raised ruminants has a higher carbon equivalent footprint than other meats or vegetarian sources of protein based on a global meta-analysis of lifecycle assessment studies.[47] Methane production by meat animals, principally ruminants, is estimated 15–20% global production of methane, unless the animals were hunted in the wild.[48][49] The current U.S. domestic beef and dairy cattle population is around 90 million head, approximately 50% higher than the peak wild population of American bison of 60 million head in the 1700s,[50] which primarily roamed the part of North America that now makes up the United States.
Artiodactyls are placental mammals belonging to the order Artiodactyla. Typically, they are ungulates which bear weight equally on two of their five toes. The other three toes are either present, absent, vestigial, or pointing posteriorly. By contrast, most perissodactyls bear weight on an odd number of the five toes. Another difference between the two orders is that many artiodactyls digest plant cellulose in one or more stomach chambers rather than in their intestine. Molecular biology, along with new fossil discoveries, has found that cetaceans fall within this taxonomic branch, being most closely related to hippopotamuses. Some modern taxonomists thus apply the name Cetartiodactyla to this group, while others opt to include cetaceans within the existing name of Artiodactyla. Some researchers use "even-toed ungulates" to exclude cetaceans and only include terrestrial artiodactyls, making the term paraphyletic in nature.
Dietary fiber or roughage is the portion of plant-derived food that cannot be completely broken down by human digestive enzymes. Dietary fibers are diverse in chemical composition and can be grouped generally by their solubility, viscosity and fermentability which affect how fibers are processed in the body. Dietary fiber has two main subtypes: soluble fiber and insoluble fiber which are components of plant-based foods such as legumes, whole grains, cereals, vegetables, fruits, and nuts or seeds. A diet high in regular fiber consumption is generally associated with supporting health and lowering the risk of several diseases. Dietary fiber consists of non-starch polysaccharides and other plant components such as cellulose, resistant starch, resistant dextrins, inulins, lignins, chitins, pectins, beta-glucans, and oligosaccharides.
Digestion is the breakdown of large insoluble food compounds into small water-soluble components so that they can be absorbed into the blood plasma. In certain organisms, these smaller substances are absorbed through the small intestine into the blood stream. Digestion is a form of catabolism that is often divided into two processes based on how food is broken down: mechanical and chemical digestion. The term mechanical digestion refers to the physical breakdown of large pieces of food into smaller pieces which can subsequently be accessed by digestive enzymes. Mechanical digestion takes place in the mouth through mastication and in the small intestine through segmentation contractions. In chemical digestion, enzymes break down food into the small compounds that the body can use.
The Bovidae comprise the biological family of cloven-hoofed, ruminant mammals that includes cattle, bison, buffalo, antelopes, sheep and goats. A member of this family is called a bovid. With 143 extant species and 300 known extinct species, the family Bovidae consists of 11 major subfamilies and thirteen major tribes. The family evolved 20 million years ago, in the early Miocene.
Chewing or mastication is the process by which food is crushed and ground by the teeth. It is the first step in the process of digestion, allowing a greater surface area for digestive enzymes to break down the foods.
Cud is a portion of food that returns from a ruminant's stomach to the mouth to be chewed for the second time. More precisely, it is a bolus of semi-degraded food regurgitated from the reticulorumen of a ruminant. Cud is produced during the physical digestive process of rumination.
A monogastric organism has a simple single-chambered stomach. Examples of monogastric omnivores include humans, pigs, hamsters and rats. Furthermore, there are monogastric carnivores such as cats. A monogastric organism is contrasted with ruminant organisms, such as cattle, goats, and sheep. Herbivores with monogastric digestion can digest cellulose in their diets by way of symbiotic gut bacteria. However, their ability to extract energy from cellulose digestion is less efficient than in ruminants.
The rumen, also known as a paunch, is the largest stomach compartment in ruminants. The rumen and the reticulum make up the reticulorumen in ruminant animals. The diverse microbial communities in the rumen allows it to serve as the primary site for microbial fermentation of ingested feed, which is often fiber-rich roughage typically indigestible by mammalian digestive systems. The rumen is known for containing unique microbial networks within its multiple sac compartments to break down nutrients into usable energy and fatty acids.
The omasum, also known as the bible, the fardel, the manyplies and the psalterium, is the third compartment of the stomach in ruminants. The omasum comes after the rumen and reticulum and before the abomasum. Different ruminants have different omasum structures and function based on the food that they eat and how they developed through evolution.
Enteric fermentation is a digestive process by which carbohydrates are broken down by microorganisms into simple molecules for absorption into the bloodstream of an animal. Because of human agricultural reliance in many parts of the world on animals which digest by enteric fermentation, it is the second largest anthropogenic factor for the increase in methane emissions directly after fossil fuel use.
The abomasum, also known as the maw, rennet-bag, or reed tripe, is the fourth and final stomach compartment in ruminants. It secretes rennet, which is used in cheese creation.
Cecotropes are a nutrient filled package created in the gastointestinal (GI) tract, expelled and eaten by rabbits and guinea pigs to get more nutrition out of their food. The first time through the GI tract, small particles of fiber are moved into the cecum, where microbes ferment them. This creates useable nutrients which are stored and expelled in cecotropes. The cecotropes are eaten and the nutrients are absorbed in the small intestine.
Pecora is an infraorder of even-toed hoofed mammals with ruminant digestion. Most members of Pecora have cranial appendages projecting from their frontal bones; only two extant genera lack them, Hydropotes and Moschus. The name "Pecora" comes from the Latin word pecus, which means "cattle". Although most pecorans have cranial appendages, only some of these are properly called "horns", and many scientists agree that these appendages did not arise from a common ancestor, but instead evolved independently on at least two occasions. Likewise, while Pecora as a group is supported by both molecular and morphological studies, morphological support for interrelationships between pecoran families is disputed.
The Cetruminantia are a clade made up of the Cetancodontamorpha and their closest living relatives, the Ruminantia.
Fibrobacter succinogenes is a cellulolytic bacterium species in the genus Fibrobacter. It is present in the rumen of cattle. F. succinogenes is a gram negative, rod-shaped, obligate anaerobe that is a major contributor to cellulose digestion. Since its discovery in the 1950s, it has been studied for its role in herbivore digestion and cellulose fermentation, which can be utilized in biofuel production.
Hindgut fermentation is a digestive process seen in monogastric herbivores. Cellulose is digested with the aid of symbiotic microbes including bacteria, archaea, and eukaryotes. The microbial fermentation occurs in the digestive organs that follow the small intestine: the cecum and large intestine. Examples of hindgut fermenters include proboscideans and large odd-toed ungulates such as horses and rhinos, as well as small animals such as rodents, rabbits and koalas.
Ruminal tympany, also known as ruminal bloat, is a disease of ruminant animals, characterized by an excessive volume of gas in the rumen. Ruminal tympany may be primary, known as frothy bloat, or secondary, known as free-gas bloat.
Methanogens are a group of microorganisms that produce methane as a byproduct of their metabolism. They play an important role in the digestive system of ruminants. The digestive tract of ruminants contains four major parts: rumen, reticulum, omasum and abomasum. The food with saliva first passes to the rumen for breaking into smaller particles and then moves to the reticulum, where the food is broken into further smaller particles. Any indigestible particles are sent back to the rumen for rechewing. The majority of anaerobic microbes assisting the cellulose breakdown occupy the rumen and initiate the fermentation process. The animal absorbs the fatty acids, vitamins and nutrient content on passing the partially digested food from the rumen to the omasum. This decreases the pH level and initiates the release of enzymes for further breakdown of the food which later passes to the abomasum to absorb remaining nutrients before excretion. This process takes about 9–12 hours.
Displaced abomasum in cattle occurs when the abomasum, also known as the true stomach, which typically resides on the floor of the abdomen, fills with gas and rises to the top of the abdomen, where it is said to be ‘displaced’. When the abomasum moves from its normal position it prevents the natural passage of gas and feed through the digestive system, creating a restriction. As cattle are ruminants, which have a 4 chambered stomach composed of a rumen, reticulum, omasum and abomasum. Ruminants require this specialized digestive system in order to properly process and break down their high fiber and cellulose rich diets. As this type of digestive system is quite complex it is at a greater risk for incidence. Due to the natural anatomy of cattle it is more common to have the abomasum displace to the left, known as a left-displaced abomasum, than to the right, right-displaced abomasum. When the abomasum becomes displaced there also becomes a chance of an abomasal volvulus, twist, developing. An abomasal volvulus occurs when the abomasum, which is already out of place, will rotate and cut off blood and nutrient supply to the abomasum. Cattle which develop an abomasal twist require immediate vet attention to regain blood supply and food passage through the digestive system or the abomasum will begin to shut down due to lack of blood supply and toxicity development.
The Jarman–Bell principle is a concept in ecology that the food quality of a herbivore's intake decreases as the size of the herbivore increases, but the amount of such food increases to counteract the low quality foods. It operates by observing the allometric properties of herbivores. The principle was coined by P.J Jarman (1968.) and R.H.V Bell (1971).
1 2 3 Fernández, Manuel Hernández; Vrba, Elisabeth S. (1 May 2005). "A complete estimate of the phylogenetic relationships in Ruminantia: a dated species-level supertree of the extant ruminants". Biological Reviews. 80 (2): 269–302. doi:10.1017/s1464793104006670. ISSN1469-185X. PMID15921052. S2CID29939520.
1 2 Fowler, M.E. (2010). "Medicine and Surgery of Camelids", Ames, Iowa: Wiley-Blackwell. Chapter 1 General Biology and Evolution addresses the fact that camelids (including camels and llamas) are not ruminants, pseudo-ruminants, or modified ruminants.
↑ Austin, PJ; etal. (1989). "Tannin-binding proteins in saliva of deer and their absence in saliva of sheep and cattle". J Chem Ecol. 15 (4): 1335–47. doi:10.1007/BF01014834. PMID24272016. S2CID32846214.
↑ Ripple, William J.; Pete Smith; Helmut Haberl; Stephen A. Montzka; Clive McAlpine & Douglas H. Boucher. 2014. "Ruminants, climate change and climate policy". Nature Climate Change. Volume 4 No. 1. pp. 2–5.
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