Hindgut fermentation
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Hindgut fermentation is a digestive process seen in
monogastric A monogastric organism has a simple single-chambered stomach (one stomach). Examples of monogastric herbivores are horses and rabbits. Examples of monogastric omnivores include humans, pigs, hamsters and rats. Furthermore, there are monogastric ...
herbivores, animals with a simple, single-chambered
stomach The stomach is a muscular, hollow organ in the gastrointestinal tract of humans and many other animals, including several invertebrates. The stomach has a dilated structure and functions as a vital organ in the digestive system. The stomach i ...
.
Cellulose Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell w ...
is digested with the aid of symbiotic
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
.Animal Structure & Function
. Sci.waikato.ac.nz. Retrieved on 2011-11-27.
The microbial fermentation occurs in the digestive organs that follow the small intestine: the
large intestine The large intestine, also known as the large bowel, is the last part of the gastrointestinal tract and of the digestive system in tetrapods. Water is absorbed here and the remaining waste material is stored in the rectum as feces before bein ...
and
cecum The cecum or caecum is a pouch within the peritoneum that is considered to be the beginning of the large intestine. It is typically located on the right side of the body (the same side of the body as the appendix, to which it is joined). The wo ...
. Examples of hindgut fermenters include proboscideans and large
odd-toed ungulate Odd-toed ungulates, mammals which constitute the taxonomic order Perissodactyla (, ), are animals—ungulates—who have reduced the weight-bearing toes to three (rhinoceroses and tapirs, with tapirs still using four toes on the front legs) o ...
s such as horses and
rhino A rhinoceros (; ; ), commonly abbreviated to rhino, is a member of any of the five extant species (or numerous extinct species) of odd-toed ungulates in the family Rhinocerotidae. (It can also refer to a member of any of the extinct species o ...
s, as well as small animals such as
rodent Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are n ...
s, rabbits and
koala The koala or, inaccurately, koala bear (''Phascolarctos cinereus''), is an arboreal herbivorous marsupial native to Australia. It is the only extant representative of the family Phascolarctidae and its closest living relatives are the w ...
s. In contrast, foregut fermentation is the form of cellulose digestion seen in
ruminant Ruminants (suborder Ruminantia) are hoofed herbivorous grazing or browsing mammals 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 ...
s such as
cattle Cattle (''Bos taurus'') are large, domesticated, cloven-hooved, herbivores. They are a prominent modern member of the subfamily Bovinae and the most widespread species of the genus ''Bos''. Adult females are referred to as cows and adult ma ...
which have a four-chambered stomach, as well as in
sloth Sloths are a group of Neotropical xenarthran mammals constituting the suborder Folivora, including the extant arboreal tree sloths and extinct terrestrial ground sloths. Noted for their slowness of movement, tree sloths spend most of their l ...
s,
macropodid Macropodidae is a family of marsupials that includes kangaroos, wallabies, tree-kangaroos, wallaroos, pademelons, quokkas, and several other groups. These genera are allied to the suborder Macropodiformes, containing other macropods, and ar ...
s, some monkeys, and one bird, the
hoatzin The hoatzin ( ) or hoactzin ( ), (''Opisthocomus hoazin''), is the only species in the order Opisthocomiformes. It is a species of tropical bird found in swamps, riparian forests, and mangroves of the Amazon and the Orinoco basins in South Ameri ...
..


Cecum

Hindgut fermenters generally have a cecum and large intestine that are much larger and more complex than those of a foregut or
midgut The midgut is the portion of the embryo from which most of the intestines develop. After it bends around the superior mesenteric artery, it is called the "midgut loop". It comprises the portion of the alimentary canal from the end of the foregut a ...
fermenter. Research on small cecum fermenters such as flying squirrels, rabbits and lemurs has revealed these mammals to have a GI tract about 10-13 times the length of their body. This is due to the high intake of
fiber Fiber or fibre (from la, fibra, links=no) is a natural or artificial substance that is significantly longer than it is wide. Fibers are often used in the manufacture of other materials. The strongest engineering materials often incorpora ...
and other hard to digest compounds that are characteristic to the diet of monogastric herbivores. Unlike in foregut fermenters, the cecum is located after the
stomach The stomach is a muscular, hollow organ in the gastrointestinal tract of humans and many other animals, including several invertebrates. The stomach has a dilated structure and functions as a vital organ in the digestive system. The stomach i ...
and small intestine in monogastric animals, which limits the amount of further digestion or absorption that can occur after the food is fermented.


Large intestine

In smaller hindgut fermenters of the order
Lagomorpha The lagomorphs are the members of the taxonomic order Lagomorpha, of which there are two living families: the Leporidae (hares and rabbits) and the Ochotonidae ( pikas). The name of the order is derived from the Ancient Greek ''lagos'' (λαγ ...
(rabbits, hares, and
pika A pika ( or ; archaically spelled pica) is a small, mountain-dwelling mammal found in Asia and North America. With short limbs, very round body, an even coat of fur, and no external tail, they resemble their close relative, the rabbit, but wi ...
s), cecotropes formed in the cecum are passed through the large intestine and subsequently reingested to allow another opportunity to absorb nutrients. Cecotropes are surrounded by a layer of mucus which protects them from stomach acid but which does not inhibit nutrient absorption in the small intestine.
Coprophagy Coprophagia () or coprophagy () is the consumption of feces. The word is derived from the grc, κόπρος , "feces" and , "to eat". Coprophagy refers to many kinds of feces-eating, including eating feces of other species (heterospecifics), o ...
is also practiced by some rodents, such as the capybara,
guinea pig The guinea pig or domestic guinea pig (''Cavia porcellus''), also known as the cavy or domestic cavy (), is a species of rodent belonging to the genus '' Cavia'' in the family Caviidae. Breeders tend to use the word ''cavy'' to describe the ...
and related species, and by the marsupial common ringtail possum. This process is also beneficial in allowing for restoration of the microflora population, or gut flora. These microbes are found in the digestive organs of living creatures and can act as protective agents that strengthen the
immune system The immune system is a network of biological processes that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to parasitic worms, as well as cancer cells and objects such as wood splint ...
. Hindgut fermenters do have the ability to expel their microflora, which is useful during the acts of hibernation,
estivation Aestivation ( la, aestas (summer); also spelled estivation in American English) is a state of animal dormancy, similar to hibernation, although taking place in the summer rather than the winter. Aestivation is characterized by inactivity and a ...
and
torpor Torpor is a state of decreased physiological activity in an animal, usually marked by a reduced body temperature and metabolic rate. Torpor enables animals to survive periods of reduced food availability. The term "torpor" can refer to the time ...
.


Efficiency

While foregut fermentation is generally considered more efficient, and monogastric animals cannot digest cellulose as efficiently as ruminants, hindgut fermentation allows animals to consume small amounts of low-quality forage all day long and thus survive in conditions where ruminants might not be able to obtain nutrition adequate for their needs. Hindgut fermenters are able to extract more nutrition out of small quantities of feed. The large hind-gut fermenters are bulk feeders: they ingest large quantities of low-nutrient food, which they process more rapidly than would be possible for a similarly sized foregut fermenter. The main food in that category is grass, and grassland grazers move over long distances to take advantage of the growth phases of grass in different regions.


Speed

The ability to process food more rapidly than foregut fermenters gives hindgut fermenters an advantage at very large body size, as they are able to accommodate significantly larger food intakes. The largest extant and prehistoric megaherbivores, elephants and indricotheres (a type of rhino), respectively, have been hindgut fermenters. Study of the rates of evolution of larger maximum body mass in different terrestrial mammalian groups has shown that the fastest growth in body mass over time occurred in hindgut fermenters (
perissodactyl Odd-toed ungulates, mammals which constitute the taxonomic order Perissodactyla (, ), are animals—ungulates—who have reduced the weight-bearing toes to three (rhinoceroses and tapirs, with tapirs still using four toes on the front legs) o ...
s,
rodent Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are n ...
s and proboscids).


Types

Hindgut fermenters are subdivided into two groups based on the relative size of various digestive organs in relationship to the rest of the system: colonic fermenters tend to be larger species such as horses, and cecal fermenters are smaller animals such as rabbits and rodents.Grant, Kerri
Adaptations in Herbivore Nutrition
July 30, 2010. Lafebervet.com. Retrieved on 2017-10-16.
However, in spite of the terminology, colonic fermenters such as horses make extensive use of the cecum to break down cellulose. Also, colonic fermenters typically have a proportionally longer large intestine than small intestine, whereas cecal fermenters have a considerably enlarged cecum compared to the rest of the digestive tract.


Insects

In addition to mammals, several insects are also hindgut fermenters, the best studied of which are the
termites Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blattod ...
, which are characterised by an enlarged "paunch" of the hindgut that also houses the bulk of the gut microbiota. Digestion of wood particles in lower termites is accomplished inside the phagosomes of gut flagellates, but in the flagellate-free higher termites, this appears to be accomplished by fibre-associated bacteria.


See also

* Foregut fermentation *
Pseudoruminant Pseudoruminant is a classification of animals based on their digestive tract differing from the ruminants. Hippopotami and camels are ungulate mammals with a three-chambered stomach (ruminants have a four-chambered stomach) while equids (horses, ...
s *
Ruminant Ruminants (suborder Ruminantia) are hoofed herbivorous grazing or browsing mammals 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 ...
s


References

{{reflist, 2 Digestive system Biology terminology