Are Horses Ruminant Mammals

Elaine Sutton
• Wednesday, 21 October, 2020
• 41 min read

Since many animals that survive on grassy forage tend to be ruminants, does that mean that horses are too? Ruminant animals have four compartments within their stomach that digest their food in stages.

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Horses only have one compartment in their stomach which means they do not fall within the ruminant category. It is a common misconception that horses are ruminant animals like cows.

Animals that are ruminants have stomachs that are divided into four compartments, each of which performs a different function. Lots of microorganisms live within the rumen, and they work hard to help break down the forage that the animal eats.

When a ruminant animal eats some grass or other forage, they do not entirely chew up the food. This cud is then pushed back up to the animal’s mouth, rec hewed and then swallowed again.

This helps the animal’s digestive system break down the cellulose that comprises the grass or forage they have eaten. Examples of Ruminant AnimalsExamples often- Ruminant Animals Cows Horses Deer Pigs Sheep Chickens Goats Dogs Moose Cats Elk Lions Bison Chimpanzees A horse has a stomach that contains only one compartment.

A horse’s stomach and small intestine function much like other monogastric animals like dogs, cats and pigs. It is able to process cellulose, a substrate found in grass and vegetation that is impossible for humans to digest.

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It is basically a fermentation container that works to break down the forage, specifically the cellulose, that horses eat. So, in simple terms, a horse chews its food completely the first time and swallows it.

Instead of using the lumen to process cellulose (a plant substance that is non-digestible for humans) horses can use their large intestines, specifically the cecum, to perform this function. So, in a way, the cecum performs the same job as the lumen in ruminant animals.

Cows, well-known ruminants, chew their food in systematic, rhythmic ways. Horses cannot rec hew their cud like ruminant animals do, but they do chew their food extensively in an effort to prepare it for the fermentation it will encounter in the upper part of their large intestine.

Many people make the mistake of thinking that some animals, like cattle, have 4 separate stomachs. Some animals, like ruminants, have multiple compartments within their single stomach.

The C-1 part of the stomach is most similar to the lumen compartment in a ruminant animal. Many people mistakenly believe that this means that the Merychippus horse was a ruminant animal.

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The prehistoric horse Merychippus was physiologically unable to do that, and they only had one chamber in their stomach. Horses have only one chamber within their stomach compared to the four compartments that ruminant animals possess.

The process of rec hewing the cud to further break down plant matter and stimulate digestion is called rumination. The word ruminant comes from the Latin ruminate, which means “to chew over again”.

Hoffmann 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, lumen size, and lumen papillae. 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 Hoffmann and Stewart warrant further research.

However, their anatomy and method of digestion differs significantly from that of a four-chambered ruminant. A, dog; B, Mus documents ; C, Mus muscles ; D, weasel; E, scheme of the ruminant stomach, the arrow with the dotted line showing the course taken by the food; F, human stomach.

These two compartments make up the fermentation vat, they 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 utilize them.

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Microbes function best in a warm, moist, anaerobic environment with a temperature range of 37.7 to 42.2 °C (100 to 108 °F) and a pH between 6.0 and 6.4. Without the help of microbes, ruminants would not be able to utilize nutrients from forages.

The food is mixed with saliva and separates into layers of solid and liquid material. 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.

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 lumen pH.

The type of feed the animal consumes affects the amount of saliva that is produced. Though the lumen and reticulum have different names, they have very similar tissue layers and textures, making it difficult to visually separate them.

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The degraded digest, which is now in the lower liquid part of the reticulorumen, then passes into the next chamber, the oakum. It keeps the particle size as small as possible in order to pass into the aromas.

The oakum also absorbs volatile fatty acids and ammonia. The aromas is the direct equivalent of the monogastric stomach, and digest is digested here in much the same way.

This compartment releases acids and enzymes that further digest the material passing through. Digest 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 digest 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.

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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. Fermentation continues in the large intestine in the same way as in the reticulorumen.

Only small amounts of glucose are absorbed from dietary carbohydrates. 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 Via propitiate, glycerol, lactate, and protein.

The Via propitiate is used for around 70% of the glucose and glycogen produced and protein for another 20% (50% under starvation conditions). Wild ruminants number at least 75 million and are native to all continents except Antarctica.

Species inhabit a wide range of climates (from tropic to arctic) and habitats (from open plains to forests). Ruminating animals have various physiological features that enable them to survive in nature.

During grazing, the silica content in forage causes abrasion of the teeth. This abrasion 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.

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Most ruminants do not have upper incisors; instead, they have a thick dental pad to thoroughly chew plant-based food. Another feature of ruminants is the large luminal 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, rec hewing, resalivation, and res wallowing. Rumination reduces particle size, which enhances microbial function and allows the digest to pass more easily through the digestive tract.

Vertebrates lack the ability to hydrolyze the beta glycosidic bond of plant cellulose due to the lack of the enzyme cellulose. Thus, ruminants must completely depend on the microbial flora, present in the lumen or hind gut, to digest cellulose.

The hydrolysis of cellulose results in sugars, which are further fermented to acetate, lactate, propitiate, literate, carbon dioxide, and methane. As bacteria conduct fermentation in the lumen, they consume about 10% of the carbon, 60% of the phosphorus, and 80% of the nitrogen that the ruminant ingests.

To reclaim these nutrients, the ruminant then digests the bacteria in the aromas. The enzyme lysosome has adapted to facilitate digestion of bacteria in the ruminant aromas.

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Pancreatic ribonuclease also degrades bacterial RNA in the ruminant small intestine as a source of nitrogen. The role of saliva is to provide ample fluid for lumen fermentation and to act as a buffering agent.

After digest pass through the lumen, the oakum absorbs excess fluid so that digestive enzymes and acid in the aromas are not diluted. 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 species of plants. 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. 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 lumen bacteria.

Very high levels of tannin intake can produce toxicity that can even cause death. 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.

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Some ruminants (goats, deer, elk, moose) are able to consume feed high in tannins (leaves, twigs, bark) due to the presence in their saliva of tannin-binding proteins. The verb 'to ruminate' has been extended metaphorically to mean to ponder thoughtfully or to meditate on some topic.

In psychology, “rumination” refers to a pattern of thinking, and is unrelated to digestive physiology. In 2010, enteric fermentation accounted for 43% of the total greenhouse gas emissions from all agricultural activity in the world, 26% of the total greenhouse gas emissions from agricultural activity in the U.S., and 22% of the total U.S. methane emissions.

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. Methane production by meat animals, principally ruminants, is estimated 15–20% global production of methane, unless the animals were hunted in the wild.

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, which primarily roamed the part of North America that now makes up the United States. ^ a b c Fernández, Manuel Hernández; VBA, Elisabeth S. (2005-05-01).

“A complete estimate of the phylogenetic relationships in Ruminants: a dated species-level super tree of the extant ruminants”. Chapter 1 General Biology and Evolution addresses the fact that came lids (including camels and llamas) are not ruminants, pseudo-ruminants, or modified ruminants.

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^ Richard F. Kay, M. Susana Cargo, Early Miocene Paleo biology in Patagonia: High-Latitude Paleo communities of the Santa Cruz Formation, Cambridge University Press, 11/10/2012 ^ “Suborder Ruminating, the Ultimate Ungulate”. “Evolutionary steps of physiological and diversification of ruminants: a comparative view of their digestive system”.

Functional Anatomy and Physiology of Domestic Animals, pages 357–358 ISBN 978-0-7817-4333-4 ^ Colorado State University, Hypertext for Biomedical Science: Nutrient Absorption and Utilization in Ruminants ^ a b c d Hickman. Ruminant ecology and evolution: Perspectives useful to livestock research and production”.

Journal of Dairy Science, 93:1320–1334 ^ “Dental Anatomy of Ruminants”. “Reconstructing the evolutionary history of the artiodactyl ribonuclease super family” (PDF).

“Some physical and chemical properties of Bovine saliva which may affect lumen digestion and synthesis”. “Old world ruminant morphophysiology, life history, and fossil record: exploring key innovations of a diversification sequence” (PDF).

^ a b c B. R Min, et al. (2003) The effect of condensed tannins on the nutrition and health of ruminants fed fresh temperate forages: a review Animal Feed Science and Technology 106(1):3–19 ^ Bate-Smith and Swain (1962). ^ Leviticus 11:3 ^ Asama, Nariño; Minamoto, MIA; Hind, Tune (1999).

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“Effect of the Addition of Fumarate on Methane Production by Luminal Microorganisms in Vito”. 8–58 (PDF) ^ Shin dell, D. T.; Faludi, G.; Koch, D. M.; Schmidt, G. A.; Unger, N.; Bauer, S. E. (2009).

^ Shin dell, D. T.; Faludi, G.; Koch, D. M.; Schmidt, G. A.; Unger, N.; Bauer, S. E. (2009). ^ Ripple, William J.; Pete Smith; Helmut Haber; Stephen A. Montana; Clive McAlpin & Douglas H. Boucher.

In large small ruminant production systems, a percentage of animals should be tested to determine if the herd needs to be treated. Protocols will vary based on the type, location, herd, and individual animals of an operation.

Florida peak season for equine parasite infections are spring, fall, and winter, treatments should be focused during these time frames. Fecal egg counts are often a small cost through your veterinarian and often do not require a farm visit.

Mow and harrow pastures regularly to break up manure and expose parasite eggs to the sun. Prevent overgrazing and reduce fecal contamination by keeping the number of horses per acre to a minimum.

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Remove bot eggs from the hair routinely to prevent ingestion. Sheep & Goat deforming treatments should be made using Yamaha or fecal egg count (FEC) scoring.

FEC scoring, like in other livestock species, is the practice of counting parasite eggs in a fecal sample. FEC scoring can give a more specific recommendation since species of parasite can be determined, this leads to a more targeted deforming protocol.

Deforming strategies for cattle are unlike other livestock species since they are treated by management category and not by individual animal. These management categories are most susceptible to clinical affects of parasites such as poor performance or death, they also are the heaviest shredders of eggs and the reason for most re-infection.

There are products that can effectively dose reformers without having to work cattle, such as blocks, salt/mineral mix, top dress, cubes, and others. Adult cattle have a strong immunity to parasites and generally anthelminitic (parasite control) treatments do not provide a return on investment unless cattle are showing clinical signs of infection (severe weight loss/anemia).

Summer treatments, especially those with larvae inhibitors, are highly effective in reducing overall pasture contamination and fall re-infection rates. Summer treatments should use products such as Panacea at double dose (fenbendazole), Isomer (ivermectin), Ablaze (Allendale), or Synaptic (oxfendazole) as these are the approved cattle reformers that inhibit larvae development.

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Animals treated with fenbendazole or oxfendazole can return to grazing pastures 8-12 hours after treatment. Animals treated with ivermectin should not be returned to grazing pastures for at least 48 hours after treatment.

If animals treated with ivermectin are returned to grazing pastures before 48 hours have passed viable eggs will still be shed in manure and will lead to re-infection/pasture contamination. If pasture has been grazed by other cattle in the last 6 months, then retreat in 30 days to ensure effective blocking of the parasite life cycle.

Overstocked pastures will require more treatments as cattle are more likely to come in contact with parasite larvae. Liver fluke treatments Treatment for Liver flukes should focus Spring (March/April) with Curate (clorsulon) as it is effective against both mature and immature bots, and fall (October-December) with Ablaze or Isomer F as these are effective only against mature bots.

The other three toes are either present, absent, vestigial, or pointing posteriorly. The aquatic cetaceans (whales, dolphins, and porpoises) evolved from even-toed ungulates, so modern taxonomic classification combines the two under the name Cetartiodactyla.

The roughly 270 land-based even-toed ungulate species include pigs, peccaries, hippopotamuses, antelopes, mouse deer, deer, giraffes, camels, llamas, alpacas, sheep, goats, and cattle. Many of these are of great dietary, economic, and cultural importance to humans.

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The oldest fossils of even-toed ungulates date back to the early Eocene (about 53 million years ago). Since these findings almost simultaneously appeared in Europe, Asia, and North America, it is very difficult to accurately determine the origin of artiodactyls.

The fossils are classified as belonging to the family Dichobunidae ; their best-known and best-preserved member is Diacodexis. The early to middle Eocene saw the emergence of the ancestors of most of today's mammals.

Entelodonts were stocky animals with a large head, and were characterized by bony bumps on the lower jaw. Two formerly widespread, but now extinct, families of even-toed ungulates were Entelodontidae and Anthracotheriidae. Entelodonts existed from the middle Eocene to the early Miocene in Eurasia and North America.

They had a stocky body with short legs and a massive head, which was characterized by two humps on the lower jaw bone. Anthracotheres had a large, porcine (pig -like) build, with short legs and an elongated muzzle.

This group appeared in the middle Eocene up until the Pliocene, and spread throughout Eurasia, Africa, and North America. Anthracotheres are thought to be the ancestors of hippos, and, likewise, probably led a similar aquatic lifestyle.

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Among the North American camels were groups like the stocky, short-legged Merycoidodontidae. They first appeared in the late Eocene and developed a great diversity of species in North America.

Only in the late Miocene or early Pliocene did they migrate from North America into Eurasia. The North American varieties became extinct around 10,000 years ago.

In the late Eocene or the Oligocene, two families stayed in Eurasia and Africa; the peccaries, which became extinct in the Old World, exist today only in the Americas. The classification of artiodactyls was hotly debated because the ocean-dwelling cetaceans evolved from the land-dwelling even-toed ungulates.

To address this problem, the traditional order Artiodactyla and infra order Cetacea are sometimes subsumed into the more inclusive Cetartiodactyla taxon. An alternative approach is to include both land-dwelling even-toed ungulates and ocean-dwelling cetaceans in a revised Artiodactyla taxon.

Molecular and morphological studies confirmed that cetaceans are the closest living relatives of hippopotamuses. In the 1990s, biological systematic used not only morphology and fossils to classify organisms, but also molecular biology. Comparison of even-toed ungulate and cetaceans genetic material has shown that the closest living relatives of whales and hippopotamuses is the paraplegic group Artiodactyla.

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They were both archdioceses (“ancient whales”) from about 48 million years ago (in the Eocene). These findings showed that archdioceses were more terrestrial than previously thought, and that the special construction of the talus (ankle bone) with a double-rolled joint surface, previously thought to be unique to even-toed ungulates, were also in early cetaceans.

The mesonychids, another type of ungulate, did not show this special construction of the talus, and thus was concluded to not have the same ancestors as cetaceans. Hippos are a geologically young group, which raises questions about their origin. The oldest cetaceans date back to the early Eocene (53 million years ago), whereas the oldest known hippopotamus dates back only to the Miocene (15 million years ago).

Some doubts have arisen regarding the relationship between the two, as there is a 40-million-year gap between their first appearances in the fossil record. It seems unlikely that there were ancestral hippos that left no remains, given the high number of even-toed ungulate fossils.

Some studies proposed the late emergence of hippos is because they are relatives of peccaries and split recently, but molecular findings contradict this. Research is therefore focused on anthracortheres (family Anthracotheriidae); one dating from the Eocene to Miocene was declared to be “hippo-like” upon discovery in the 19th century.

A study from 2005 showed that the anthracotheres and hippopotamuses have very similar skulls, but differed in the adaptations of their teeth. It was nevertheless believed that cetaceans and anthracothereres descended from a common ancestor, and that hippopotamuses developed from anthracotheres.

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A study published in 2015 was able to confirm this, but also revealed that hippopotamuses were derived from older anthracotheriens. The newly introduced genus Epirigenys from eastern Africa is thus the sister group of hippos.

Spines (including pigs) and hippopotamuses have molars with well-developed roots and a simple stomach that digests food. All other even-toed ungulates have molars with a selenodont construction (crescent-shaped cusps) and have the ability to ruminate, which requires regurgitating food and re-chewing it.

Their molars were adapted to a carnivorous diet, resembling the teeth in modern toothed whales, and, unlike other mammals, have a uniform construction. Molecular findings and morphological indications suggest that artiodactyls as traditionally defined are paraplegic with respect to cetaceans.

Cetaceans are deeply nested within the former; the two groups together form a monophyletic taxon, for which the name Cetartiodactyla is sometimes used. Modern nomenclature divides Artiodactyla (or Cetartiodactyla) in four subordinate taxa: came lids (Toyoda), pigs and peccaries (Soon), ruminants (Ruminants), and hippos plus whales (Whippomorpha).

The peccaries (Tayassuidae) are named after glands on their belly and are indigenous to Central and South America. The ruminants (Ruminants) consist of six families: The mouse deer (Tragulidae) are the smallest and most primitive even-toed-ruminants; they inhabit forests of Africa and Asia.

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The antilocaprids (Antilocapridae) of North America comprise only one extant species: the pronghorn. The deer (Cervical) are made up of about 45 species, which are characterized by a pair of antlers (generally only in males).

Two major body types are known: Suicide and hippopotamuses are characterized by a stocky body, short legs, and a large head; camels and ruminants, though, have a more slender build and lanky legs. Size varies considerably; the smallest member, the mouse deer, often reaches a body length of only 45 centimeters (18 in) and a weight of 1.5 kilograms (3.3 lb).

All even-toed ungulates display some form of sexual dimorphism : the males are consistently larger and heavier than the females. In deer, only the males boast antlers, and the horns of bovines are usually small or not present in females.

Fur varies in length and coloration depending on the habitat. Camouflaged coats come in colors of yellow, gray, brown, or black tones.

The first toe is missing in modern artiodactyls, and can only be found in now-extinct genera. When camels have only two toes present, the claws are transformed into nails.

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These claws consist of three parts: the plate (top and sides), the sole (bottom), and the bale (rear). In general, the claws of the forelegs are wider and blunter than those of the hind legs, and the gape is farther apart.

Aside from camels, all even-toed ungulates put just the tip of the foremost phalanx on the ground. Diagrams of hand skeletons of various mammals, left to right: orangutan, dog, pig, cow, tapir, and horse.

The muscles of the limbs are predominantly localized, which ensures that artiodactyls often have very slender legs. A clavicle is never present, and the scapula is very agile and swings back and forth for added mobility when running.

In addition, many smaller artiodactyls have a very flexible body, contributing to their speed by increasing their stride length. The skull is elongated and rather narrow; the frontal bone is enlarged near the back and displaces the parietal bone, which forms only part of the side of the cranium (especially in ruminants).

True horns have a bone core that is covered in a permanent sheath of keratin, and are found only in the bodies. Antlers are bony structures that are shed and replaced each year; they are found in deer (members of the family Cervical).

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They grow from a permanent outgrowth of the frontal bone called the pedicle and can be branched, as in the white-tailed deer (Odocoileus Virginians), or palmate, as in the moose (Alces). Pronghorns, while similar to horns in that they have gelatinous sheaths covering permanent bone cores, are deciduous.

All these cranial appendages can serve for posturing, battling for mating privilege, and for defense. The Soon and hippopotamuses have a relatively large number of teeth (with some pigs having 44); their dentition is more adapted to a squeezing mastication, which is characteristic of omnivores.

The incisors are often reduced in ruminants, and are completely absent in the upper jaw. The canines are enlarged and tusk-like in the Soon, and are used for digging in the ground and for defense.

Similar to many other prey animals, their eyes are on the sides of the head, giving them an almost panoramic view. Ruminants' mouths often have additional salivary glands, and the oral mucosa is often heavily calloused to avoid injury from hard plant parts and to allow easier transport of roughly chewed food.

Their stomachs are divided into three to four sections: the lumen, the reticulum, the oakum, and the aromas. After the food is ingested, it is mixed with saliva in the lumen and reticulum and separates into layers of solid versus liquid material.

The solids lump together to form a bolus (also known as the cud); this is regurgitated by reticular contractions while the glottis is closed. When the bolus enters the mouth, the fluid is squeezed out with the tongue and re-swallowed.

The bolus is chewed slowly to completely mix it with saliva and to break it down. Ingested food passes to the “fermentation chamber” (lumen and reticulum), where it is kept in continual motion by rhythmic contractions.

The handicap of a heavy digestive system has increased selective pressure towards limbs that allow the animal to quickly escape predators. The babies, however, is a herbivore, and has extra maxillary teeth to allow for proper mastication of plant material.

Most of the fermentation occurs with the help of cellulolytic microorganisms within the cecum of the large intestine. Their fore stomach has fermentation carried out by microbes and has high levels of volatile fatty acid ; it has been proposed that their complex fore stomach is a means to slow digestive passage and increase digestive efficiency.

They consume around 68 kilograms (150 lb) of grass and other plant matter each night. They may cover distances up to 32 kilometers (20 mi) to obtain food, which they digest with the help of microbes that produce cellulose.

Their closest living relatives, the whales, are obligate carnivores. The penises of even-toed ungulates have an S-shape at rest and lie in a pocket under the skin on the belly.

The corpora caverns is only slightly developed; and an erection mainly causes this curvature to extend, which leads to an extension, but not a thickening, of the penis. In some even-toed ungulates, the penis contains a structure called the urethral process.

The number of mammary glands is variable and correlates, as in all mammals, with litter size. Pigs, which have the largest litter size of all even-toed ungulates, have two rows of teats lined from the armpit to the groin area.

Secretory glands in the skin are present in virtually all species and can be located in different places, such as in the eyes; behind the horns, the neck, or back; on the feet; or in the anal region. Generally, there is a tendency to merge into larger groups, but some live alone or in pairs.

Some species also live in harem groups, with one male, several females, and their common offspring. Most artiodactyls, such as the wildebeest, are born with hair. Generally, even-toed ungulates tend to have long gestation periods, smaller litter sizes, and more highly developed newborns.

As with many other mammals, species in temperate or polar regions have a fixed mating season, while those in tropical areas breed year-round. The newborns are precocity (born relatively mature) and come with open eyes and are hairy (except the hairless hippos).

Juvenile deer and pigs have striped or spotted coats; the pattern disappears as they grow older. The juveniles of some species spend their first weeks with their mother in a safe location, where others may be running and following the herd within a few hours or days.

The artiodactyls with the longest lifespans are the hippos, cows, and camels, which can live 40 to 50 years. Some artiodactyls, like sheep, have been domesticated for thousands of years. Artiodactyls have been hunted by primitive humans for various reasons: for meat or fur, as well as to use their bones and teeth as weapons or tools.

To date, humans have domesticated goats, sheep, cattle, camels, llamas, alpacas, and pigs. Initially, livestock was used primarily for food, but they began being used for work activities around 3000 BCE.

Clear evidence exists of antelope being used for food 2 million years ago in the Olduvai Gorge, part of the Great Rift Valley. Cro-Magnons relied heavily on reindeer for food, skins, tools, and weapons; with dropping temperatures and increased reindeer numbers at the end of the Pleistocene, they became the prey of choice.

Reindeer remains accounted for 94% of bones and teeth found in a cave above the Chou River that was inhabited around 12,500 years ago. Today, artiodactyls are kept primarily for their meat, milk, and wool, fur, or hide for clothing.

Domestic cattle, the water buffalo, the yak, and camels are used for work, as rides, or as pack animals. The aurochs has been extinct since the 17th century. The endangerment level of each even-toed ungulate is different.

Some species are misanthropic (such as the wild boar) and have spread into areas that they are not indigenous to, either having been brought as farm animals or having run away as people's pets. Some artiodactyls also benefit from the fact that their predators (e.g. the Tasmanian tiger) were severely decimated by ranchers, who saw them as competition.

Conversely, many artiodactyls have declined significantly in numbers, and some have even gone extinct, largely due to over-hunting, and, more recently, habitat destruction. Extinct species include several gazelles, the aurochs, the Malagasy hippopotamus, the blue buck, and Homburg's deer.

Two species, the Scimitar-horned onyx and Père David's deer, are extinct in the wild. Fourteen species are considered critically endangered, including the ADDX, the osprey, the Bactria camel, Przewalski's gazelle, the siege, and the pygmy hog.

^ Jessica M Theodor; Jörg Erfurt; Gregory Metals (23 October 2007). “The earliest artiodactyls: Diacodexeidae, Dichobunidae, Homacodontidae, Leptochoeridae and Raoellidae”.

“Relationships of Cetacea (Artiodactyla) Among Mammals : Increased Taxon Sampling Alters Interpretations of Key Fossils and Character Evolution”. ^ a b Montgelard, Claudine; Cathexis, Francois M.; Doddery, Emmanuel (1997).

“Phylogenetic relationships of artiodactyls and cetaceans as deduced from the comparison of monochrome b and 12S rRNA mitochondrial sequences” (PDF). “Molecular Evidence for the Inclusion of Cetaceans within the Order Artiodactyla” (PDF).

^ Gates, John; Ayasdi, Cheryl; Cronin, Mathew A.; Alexander, Peter (1996). “Evidence from milk casein genes that cetaceans are close relatives of hippopotamus artiodactyls”.

“Molecular evidence from retroposons that whales form a clade within even-toed ungulates”. ^ a b ^ Gates, John; Milinkovitch, Michel; Weddell, Victor; Stan hope, Michael (1999).

“Stability of Sadistic Relationships between Cetacea and Higher-Level Artiodactyl Taxa”. ^ Madden, Ole; Williamson, Frederik; Using, Born M.; Reason, Sulfur; DE Long, Wilfred W. (2002).

^ Amrine-Madsen, Heather; Nepali, Klaus-Peter; Wayne, Robert K.; Springer, Mark S. (2003). “A new phylogenetic marker, lipoprotein B, provides compelling evidence for Eutheria relationships”.

“A complete phylogeny of the whales, dolphins and even-toed hoofed mammals (Cetartiodactyla)”. ^ ETA Noah BEI Noway (1999) Oder Hendricks (2004) ^ Malcolm C. McKenna; Susan K. Bell (1997).

“Resolving conflict in Eutheria mammal phylogeny using phylogenetic and the multispecies coalescent model”. Dos Was, M.; Income, J.; Hasegawa, M.; Asher, R.J.; Donahue, P.C.J.

“Phylogenetic datasets provide both precision and accuracy in estimating the timescale of placental mammal phylogeny”. “Inferring the mammal tree: Species-level sets of phylogeny for questions in ecology, evolution, and conservation”.

^ Coo, P.; Hi, R.; Ding, F.; QI, D.; GAO, H.; Men, H.; You, J.; HU, S.; Zhang, H. (2007). “A complete mitochondrial genome sequence of the wild two-humped camel (Camels Bactria ferns): an evolutionary history of Cambridge”.

“The Interrelationships of Higher Ruminant Families with Special Emphasis on the Members of the Cervices”. “Humans hunted for meat 2 million years ago”.

Euarchontoglires Rodentia (Rats, guinea pigs, squirrels, beavers, chinchillas, porcupines, cabanas and relatives) Lagomorpha (Rabbits and pikes) Scandinavia (Tree shrews) Vermonter (Columns) Primates (lorises, carriers, lemurs, monkeys, apes, humans) Ruminants are hoofed mammals that have a unique digestive system that allows them to better use energy from fibrous plant material than other herbivores.

Unlike monogastric such as swine and poultry, ruminants have a digestive system designed to ferment feed stuffs and provide precursors for energy for the animal to use. Anatomy of the ruminant digestive system includes the mouth, tongue, salivary glands (producing saliva for buffering lumen pH), esophagus, four-compartment stomach (lumen, reticulum, oakum, and aromas), pancreas, gall bladder, small intestine (duodenum, jejunum, and ileum), and large intestine (cecum, colon, and rectum).

On average, cattle take from 25,000 to more than 40,000 prehensile bites to harvest forage while grazing each day. These teeth crush and grind plant material during initial chewing and rumination.

Saliva’s most important function is to buffer pH levels in the reticulum and lumen. Forage and feed mixes with saliva containing sodium, potassium, phosphate, bicarbonate, and urea when consumed, to form a bolus.

Muscle contractions and pressure differences carry these substances down the esophagus to the reticulum. Left-sided view of ruminant digestive tract. Ruminants eat rapidly, swallowing much of their feed stuffs without chewing it sufficiently (< 1.5 inches).

The esophagus functions bidirectionally in ruminants, allowing them to regurgitate their cud for further chewing, if necessary. True ruminants, such as cattle, sheep, goats, deer, and antelope, have one stomach with four compartments: the lumen, reticulum, oakum, and abomasums.

The lumen is the largest stomach compartment, holding up to 40 gallons in a mature cow. Right-sided view of ruminant digestive tract. The reticulum holds approximately 5 gallons in the mature cow.

Typically, the lumen and reticulum are considered one organ because they have similar functions and are separated only by a small muscular fold of tissue. The reticulorumen is home to a population of microorganisms (microbes or “lumen bugs”) that include bacteria, protozoa, and fungi.

These microbes ferment and break down plant cell walls into their carbohydrate fractions and produce volatile fatty acids (IFAS), such as acetate (used for fat synthesis), propitiate (used for glucose synthesis), and literate from these carbohydrates. During normal digestive tract contractions, this object can penetrate the reticulum wall and make its way to the heart, where it can lead to hardware disease.

The lumen is sometimes called the “paunch.” It is lined with papillae for nutrient absorption and divided by muscular pillars into the dorsal, ventral, caudodorsal, and caudoventral sacs. Lumen microorganisms (primarily bacteria) digest cellulose from plant cell walls, digest complex starch, synthesize protein from nonprotein nitrogen, and synthesize B vitamins and vitamin K. Lumen pH typically ranges from 6.5 to 6.8.

Gases produced in the lumen include carbon dioxide, methane, and hydrogen sulfide. It is called the “many piles” or the “butcher’s bible” in reference to the many folds or leaves that resemble pages of a book.

The aromas produces hydrochloric acid and digestive enzymes, such as pepsin (breaks down proteins), and receives digestive enzymes secreted from the pancreas, such as pancreatic lipase (breaks down fats). The chief cells in the aromas secrete mucous to protect the abdominal wall from acid damage.

The small and large intestines follow the aromas as further sites of nutrient absorption. The small intestine is a tube up to 150 feet long with a 20-gallon capacity in a mature cow.

Digest entering the small intestine mix with secretions from the pancreas and liver, which elevate the pH from 2.5 to between 7 and 8. This higher pH is needed for enzymes in the small intestine to work properly.

Bile from the gall bladder is secreted into the first section of the small intestine, the duodenum, to aid in digestion. The cecum serves little function in a ruminant, unlike its role in horses.

Immature ruminants, such as young, growing calves from birth to about 2 to 3 months of age, are functionally nonruminants. This is thought to be accomplished through mature ruminants licking calves and environmental contact with these microorganisms.

Immature ruminants must undergo reticulorumen-omasal growth, including increases in volume and muscle. As ruminants develop, the reticulorumen and oakum grow rapidly and account for increasing proportions of the total stomach area.

For instance, it is recommended immature ruminants are not allowed access to feeds containing non-protein nitrogen such as urea. Design nutritional programs for ruminants considering animal age.

Concentrate selectors have a small reticulorumen in relation to body size and selectively browse trees and shrubs. Concentrate selectors are very limited in their ability to digest the fibers and cellulose in plant cell walls.

These ruminants depend on diets of grasses and other fibrous plant material. They prefer diets of fresh grasses over legumes but can adequately manage rapidly fermenting feed stuffs.

They have a fair though limited capacity to digest cellulose in plant cell walls. This allows them to “chew their cud” to reduce particle size and improve digestibility.

Once inside the reticulorumen, forage is exposed to a unique population of microbes that begin to ferment and digest the plant cell wall components and break these components down into carbohydrates and sugars. The microbes ferment sugars to produce IFAS (acetate, propitiate, literate), methane, hydrogen sulfide, and carbon dioxide.

Coupled with routine rumination (chewing and rec hewing of the cud) that increases salivary flow, this makes for a rather stable pH environment (around 6). Typically, on a high-grain diet, there is less chewing and ruminating, which leads to less salivary production and buffering agents’ being produced.

The relative concentrations of the IFAS are also changed, with propitiate being produced in the greatest quantity, followed by acetate and literate. Take care to provide adequate forage and avoid situations that might lead to acidosis when feeding ruminants high-concentrate diets.

Acidosis is discussed in detail in Mississippi State University Extension Service Publication 2519 Beef Cattle Nutritional Disorders. In addition, energy as a nutrient in ruminant diets is discussed in detail in Mississippi State University Extension Service Publication 2504 Energy in Beef Cattle Diets.

Like other living creatures, these microbes have requirements for protein and energy to facilitate growth and reproduction. Each feed stuff (such as cottonseed meal, soybean hulls, and annual rye grass forage) has different proportions of each protein type.

Excess ammonia is absorbed via the lumen wall and converted into urea in the liver, where it returns to the blood to the saliva or is excreted by the body. Toxicity occurs when the excess ammonia overwhelms the liver’s ability to detoxify it into urea.

However, with sufficient energy, microbes use ammonia and amino acids to grow and reproduce. In the aromas, the ruminant uses UIP along with microorganisms washed out of the lumen as a protein source.

The digestive system of ruminants optimizes use of lumen microbe fermentation products. Ruminants are in a unique position of being able to use such resources that are not in demand by humans but in turn provide man with a vital food source.

One of the best ways to improve agricultural sustainability is by developing and using effective ruminant livestock grazing systems. More than 60 percent of the land area in the world is too poor or erodible for cultivation but can become productive when used for ruminant grazing.

Developing a good understanding of ruminant digestive anatomy and function can help livestock producers better plan appropriate nutritional programs and properly manage ruminant animals in various production systems. A., M. A. McCann, R. H. Watson, N. N. Pietà, C. S. Homeland, A. H. Parks, B. L. Up church, N. S. Hill, and J. H. Boston.

By Jane A. Parish, PhD, Professor and Head, North Mississippi Research and Extension Center; J. Daniel Rivera, PhD, Associate Extension/Research Professor, South Mississippi Branch Experiment Station; and Holly T. Poland, PhD, former Assistant Research/Extension Professor, Animal and Dairy Sciences. Photos of ruminant digestive system courtesy of Stephanie R. Hill, PhD, former Assistant Research Professor, Animal and Dairy Sciences.

This publication may be copied and distributed without alteration for nonprofit educational purposes provided that credit is given to the Mississippi State University Extension Service. Discrimination in university employment, programs, or activities based on race, color, ethnicity, sex, pregnancy, religion, national origin, disability, age, sexual orientation, genetic information, status as a U.S. veteran, or any other status protected by applicable law is prohibited.

Extension Service of Mississippi State University, cooperating with U.S. Department of Agriculture. In today’s highly specialized world, animal nutritionists tend to consider themselves specialists in either ruminant or nonruminant nutrition.

Ruminant nutritionists appreciate the significance fermentation plays in meeting the nutritional components of animals with multiple stomachs, but they tend to think of most nonruminants as “simple-gutted.” Non ruminant nutritionists, conversely, often dismiss the importance of microbial fermentation to the health and well-being of the animal. These are critical oversights, because fermentation plays a key role in the nutritional ecology of almost every species of animal including horses.

Therefore, a basic understanding of the function fermentation plays in a wide range of species is critical when considering its importance in the horse. In terms of equine feed management, sources of starch are usually cereal grains such as oats, barley, and corn.

Animals can be divided into four basic groups according to the structure of their gastrointestinal anatomy and its ability to ferment feed stuffs. Large, multi compartment stomachs selectively sort and retain plant fiber for extended periods of time.

Hind gut fermentors are also split into two classifications according to whether they depend primarily on the cecum or colon for microbial digestion. Large nonruminant herbivores such as horses, rhinoceroses, gorillas, and elephants depend more on the colon for microbial fermentation.

Omnivores such as pigs and man have calculated colons where a good deal of digestion takes place. Non ruminant herbivores such as horses tend to dedicate a smaller proportion of their total digestive capacity to fermentation.

Both ruminant and nonruminant grazers such as horses and cows usually have more developed digestive tracts than selective herbivores like rabbits and hamsters. For example, pigs have a voluminous hind gut accounting for about 48% of their total digestive capacity, but humans devote only approximately 17% of their tracts to microbial fermentation.

As mentioned previously, carnivores usually have unsaturated colons that represent a small proportion of total digestive capacity. Retention time The extent to which plant material is fermented depends on how long it is in contact with the microbes.

Longer retention results in more complete digestion, but there is a limit to the total amount of time the material can be subjected to fermentation before energy production becomes compromised. Herbivores such as horses depend to a large degree on volatile fatty acids (IFAS) as a source of dietary energy.

If digest is retained too long in the fermenting organs, IFAS will be degraded by certain anaerobic microorganisms, thus depriving horses of energy. Animals larger than 2200 pounds must therefore employ a digestive system that is different from the ruminant to allow for rapid digest transit, which in turn supports optimal microbial fermentation.

Elephants and rhinoceroses are hind gut fermentors with digest transit times that are much faster than ruminants. These massive mammals have adopted the dietary strategy of ingesting large quantities of dry matter and passing it through the digestive system fairly quickly.

A notable exception to the relationship between body size and transit rate in nonruminants is the giant panda. They have simple, short digestive tracts with little volume to accommodate microbial fermentation, yet they live in the wild as herbivores.

The pandas have adopted a dietary strategy of extremely high intake and short retention time. Horses and elephants illustrate the general trend in rate of passage and digestibility in large nonruminant herbivores as it relates to body size.

Elephants, conversely, have a shorter retention time, about 24 hours, and lower dry matter digestibility. Despite the fact that pigs, dogs, and ponies vary tremendously in their dependence on microbial digestion, they all have hind gut environments conducive to fermentation.

Pigs are quite capable of utilizing high-fiber diets, though this fact has been largely ignored as intensive swine management programs have developed. A close look at starch, which is abundant in cereal grains such as corn, barley, and oats, proves that it is a versatile energy source.

Studies at Kentucky Equine Research (KER) have shown that pH of the hind gut drops significantly in horses following a grain meal rich in starch, with the lowest point occurring between four and eight hours after feeding. For horses that must consume large quantities of grain in order to fuel exercise or maintain body weight, a hind gut buffer is appropriate because it steadies the pH, preventing sudden downward shifts that could harm microflora.

Early work with ruminants showed that yeast culture affected microbial fermentation in a number of beneficial ways. Initially, nonruminant nutritionists dismissed this information as unimportant for monogastric animals primarily because the lumen was deemed an inappropriate model for rabbits, pigs, or horses.

The anatomical adaptations that each species has developed depend primarily on body size and natural diet. Non- ruminant animals have a simple stomach or monogastric digestive system.

In contrast, ruminant animals have a poly gastric digestive system, generally having a four-chambered stomach. Pseudo ruminants are animals that utilize large amounts of roughage or fiber as well as grains and other concentrated feeds.

Some examples of pseudo ruminants are horses, camels, alpacas, hippopotamus, rabbits, guinea pigs, and hamsters. The cecum of pseudo ruminants contains many microorganisms needed for digestion of a large amount of plants materials they consume.

Ruminants are the animals that have a poly gastric digestive system comprising a four-chambered or a multi-chambered stomach. They generally eat a large amount of roughage or fiber.

Thus, ruminants have a large stomach which has four compartments: lumen, reticulum, oakum, and aromas. Instead, they swallow their food in very large amounts, facilitating very minute chewing process.

The four compartments contain microorganisms which play a very important role in the digestion of cellulose. These microbes, especially lumen and reticulum bacteria, break down cellulose and ferment ingested food.

The oakum and the reticulum participate mainly in the grinding process of the food. The pseudo ruminant digestive system has a stomach with three compartments.

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