The vertebrate gastrointestinal tract is populated by bacteria and, in some species, protozoa and fungi that can convert dietary and endogenous substrates into absorbable nutrients. Because of a neutral pH and longer digesta retention time, the largest bacterial populations are found in the hindgut or large intestine of mammals, birds, reptiles, and adult amphibians and in the foregut of a few mammals and at least one species of bird. Bacteria ferment carbohydrates into short-chain fatty acids (SCFA), convert dietary and endogenous nitrogenous compounds into ammonia and microbial protein, and synthesize B vitamins. Absorption of SCFA provides energy for the gut epithelial cells and plays an important role in the absorption of Na and water. Ammonia absorption aids in the conservation of nitrogen and water. A larger gut capacity and longer digesta retention time provide herbivores with additional SCFA for maintenance energy and foregut-fermenting and copoprophagic hindgut-fermenting species with access to microbially synthesized protein and B vitamins. Protozoa and fungi also contribute nutrients to the host. This review discusses the contributions of gut microorganisms common to all vertebrates, the numerous digestive strategies that allow herbivores to maximize these contributions, and the effects of low-fiber diets and discontinuous feeding schedules on these microbial digestive processes.
Comparative physiology applies methods established in domestic animal science to a wider variety of species. This can lead to improved insight into evolutionary adaptations of domestic animals, by putting domestic species into a broader context. Examples include the variety of responses to seasonally fluctuating environments, different adaptations to heat and drought, and in particular adaptations to herbivory and various herbivore niches. Herbivores generally face the challenge that a high food intake compromises digestive efficiency (by reducing ingesta retention time and time available for selective feeding and for food comminution), and a variety of digestive strategies have evolved in response. Ruminants are very successful herbivores. They benefit from potential advantages of a forestomach without being constrained in their food intake as much as other foregut fermenters, because of their peculiar reticuloruminal sorting mechanism that retains food requiring further digestion but clears the forestomach of already digested material; the same mechanism also optimises food comminution. Wild ruminants vary widely in the degree to which their rumen contents 'stratify', with little stratification in 'moose-type' ruminants (which are mostly restricted to a browse niche) and a high degree of stratification into gas, particle and fluid layers in 'cattle-type' ruminants (which are more flexible as intermediate feeders and grazers). Yet all ruminants uniformly achieve efficient selective particle retention, suggesting that functions other than particle retention played an important role in the evolution of stratification-enhancing adaptations. One interesting emerging hypothesis is that the high fluid turnover observed in 'cattle-type' ruminants -which is a prerequisite for stratification -is an adaptation that not only leads to a shift of the sorting mechanism from the reticulum to the whole reticulorumen, but also optimises the harvest of microbial protein from the forestomach. Although potential benefits of this adaptation have not been quantified, the evidence for convergent evolution toward stratification suggests that they must be substantial. In modern production systems, the main way in which humans influence the efficiency of energy uptake is by manipulating diet quality. Selective breeding for conversion efficiency has resulted in notable differences between wild and domestic animals. With increased knowledge on the relevance of individual factors, that is fluid throughput through the reticulo-rumen, more specific selection parameters for breeding could be defined to increase productivity of domestic ruminants by continuing certain evolutionary trajectories.Keywords: herbivory, hindgut fermenter, foregut fermenter, browser, grazer ImplicationsUnderstanding evolutionary adaptations of ruminants will have an impact on (i) husbandry of captive wild ruminants, many of which cannot be kept or fed as domestic ruminants and (ii) research for continuous refinement of the production potential of domestic ruminants, ...
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