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Hindgut fermentation is a digestive process seen in monogastric
herbivore A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat ...
s (animals with a simple, single-chambered
stomach The stomach is a muscular, hollow organ in the upper gastrointestinal tract of Human, humans and many other animals, including several invertebrates. The Ancient Greek name for the stomach is ''gaster'' which is used as ''gastric'' in medical t ...
).
Cellulose Cellulose is an organic compound with the chemical formula, formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of glycosidic bond, β(1→4) linked glucose, D-glucose units. Cellulose is an important s ...
is digested with the aid of
symbiotic Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biolo ...
microbes including
bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...
,
archaea Archaea ( ) is a Domain (biology), domain of organisms. Traditionally, Archaea only included its Prokaryote, prokaryotic members, but this has since been found to be paraphyletic, as eukaryotes are known to have evolved from archaea. Even thou ...
, and
eukaryotes The eukaryotes ( ) constitute the domain of Eukaryota or Eukarya, organisms whose cells have a membrane-bound nucleus. All animals, plants, fungi, seaweeds, and many unicellular organisms are eukaryotes. They constitute a major group of ...
. The microbial
fermentation Fermentation is a type of anaerobic metabolism which harnesses the redox potential of the reactants to make adenosine triphosphate (ATP) and organic end products. Organic molecules, such as glucose or other sugars, are catabolized and reduce ...
occurs in the digestive organs that follow the
small intestine The small intestine or small bowel is an organ (anatomy), organ in the human gastrointestinal tract, gastrointestinal tract where most of the #Absorption, absorption of nutrients from food takes place. It lies between the stomach and large intes ...
: the
cecum The cecum ( caecum, ; plural ceca or caeca, ) 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 (a ...
and
large intestine The large intestine, also known as the large bowel, is the last part of the gastrointestinal tract and of the Digestion, digestive system in tetrapods. Water is absorbed here and the remaining waste material is stored in the rectum as feces befor ...
. Examples of hindgut fermenters include
proboscidea Proboscidea (; , ) is a taxonomic order of afrotherian mammals containing one living family (Elephantidae) and several extinct families. First described by J. Illiger in 1811, it encompasses the elephants and their close relatives. Three l ...
ns and large odd-toed ungulates such as
horses The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 milli ...
and rhinos, as well as small animals such as
rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
s, rabbits and
koala The koala (''Phascolarctos cinereus''), sometimes inaccurately called the koala bear, is an arboreal herbivorous marsupial native to Australia. It is the only Extant taxon, extant representative of the Family (biology), family ''Phascolar ...
s. In contrast, foregut fermentation is the form of cellulose digestion seen in
ruminant Ruminants are herbivorous grazing or browsing artiodactyls belonging to the suborder Ruminantia that are able to acquire nutrients from plant-based food by fermenting it in a specialized stomach prior to digestion, principally through microb ...
s such as
cattle Cattle (''Bos taurus'') are large, domesticated, bovid ungulates widely kept as livestock. They are prominent modern members of the subfamily Bovinae and the most widespread species of the genus '' Bos''. Mature female cattle are calle ...
which have a four-chambered stomach, as well as in
sloth Sloths are a Neotropical realm, Neotropical group of xenarthran mammals constituting the suborder Folivora, including the extant Arboreal locomotion, arboreal tree sloths and extinct terrestrial ground sloths. Noted for their slowness of move ...
s, macropodids, some
monkey Monkey is a common name that may refer to most mammals of the infraorder Simiiformes, also known as simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes. Thus monkeys, in that sense, co ...
s, and one bird, the hoatzin..


Cecum

Hindgut fermenters generally have a
cecum The cecum ( caecum, ; plural ceca or caeca, ) 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 (a ...
and large intestine that are much larger and more complex than those of a foregut or
midgut The midgut is the portion of the human embryo from which almost all of the small intestine and approximately half of the large intestine develop. After it bends around the superior mesenteric artery, it is called the "midgut loop". It comprises ...
fermenter.Animal Structure & Function
. Sci.waikato.ac.nz. Retrieved on 2011-11-27.
Research on small cecum fermenters such as
flying squirrel Flying squirrels (scientifically known as Pteromyini or Petauristini) are a tribe (biology), tribe of 50 species of squirrels in the family (biology), family Squirrel, Sciuridae. Despite their name, they are not in fact capable of full flight i ...
s, 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 (spelled fibre in British English; from ) 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 inco ...
and other hard to digest compounds that are characteristic to the diet of monogastric herbivores. Easily digestible food is processed in the gastrointestinal tract & expelled as regular feces. But in order to get nutrients out of hard to digest fiber, some smaller hindgut fermenters, like lagomorphs (rabbits, hares, pikas), ferment fiber in the cecum (at the small and large intestine junction) and then expel the contents as cecotropes, which are reingested ( cecotrophy). The cecotropes are then absorbed in the small intestine to utilize the nutrients. This process is also beneficial in allowing for restoration of the microflora population, or
gut flora Gut microbiota, gut microbiome, or gut flora are the microorganisms, including bacteria, archaea, fungi, and viruses, that live in the digestive tracts of animals. The gastrointestinal metagenome is the aggregate of all the genomes of the g ...
. These microbes are found in the gastrointestinal tract and can act as protective agents that strengthen the
immune system The immune system is a network of biological systems that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to bacteria, as well as Tumor immunology, cancer cells, Parasitic worm, parasitic ...
. Small hindgut fermenters have the ability to expel their microflora, which is useful during the acts of
hibernation Hibernation is a state of minimal activity and metabolic reduction entered by some animal species. Hibernation is a seasonal heterothermy characterized by low body-temperature, slow breathing and heart-rate, and low metabolic rate. It is mos ...
, estivation 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 ti ...
.


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. While ruminants require a good deal of time resting between meals, hindgut fermenters are able to take in smaller meals more frequently, allowing them to eat and move more readily. The large hindgut 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 Perissodactyla (, ), or odd-toed ungulates, is an order of Ungulate, ungulates. The order includes about 17 living species divided into three Family (biology), families: Equidae (wild horse, horses, Asinus, asses, and zebras), Rhinocerotidae ( ...
s,
rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
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 a group of detritophagous eusocial cockroaches which consume a variety of decaying plant material, generally in the form of wood, leaf litter, and soil humus. They are distinguished by their moniliform antennae and the sof ...
, 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 * Pseudoruminants *
Ruminant Ruminants are herbivorous grazing or browsing artiodactyls belonging to the suborder Ruminantia that are able to acquire nutrients from plant-based food by fermenting it in a specialized stomach prior to digestion, principally through microb ...
s * Cecotrope


References

{{reflist, 2 Digestive system Biology terminology