Evolutionary adaptations of ruminants and their potential relevance for modern production systems.

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 reticulo-rumen, 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.

Nitrogen requirement of the omnivorous greater bilby, Macrotis lagotis (Marsupialia: Peramelidae).

Mainly due to their utilisation of relatively low-fiber diets compared to herbivorous mammals, omnivores are expected to have correspondingly low maintenance nitrogen requirements (MNRs). The limited studies examining nitrogen requirements of omnivorous mammals to date have shown this to be the case. In this article, we determine the dietary MNR of greater bilbies (Macrotis lagotis), arid-zone omnivorous marsupials, by feeding them varying proportions of mixed seeds and dried currants (sun-dried grapes). We also examine the possibility that bilbies conserve nitrogen by recycling endogenous urea to their gastrointestinal tract. The dietary MNR of 127 mg N kg(-0.75) d(-1) calculated for the bilby falls within the range calculated for other marsupial omnivores and is lower than that of any of the herbivores. This low requirement for nitrogen was correlated with significant recycling of endogenous urea to the gut; bilbies recycled between 44% and 80% of urea synthesised in the liver, but the proportion recycled was independent of dietary nitrogen intake. The relatively low MNR of the bilby is consistent with its low rates of basal metabolism and the recycling of endogenous urea to the gut, both of which reduce urinary nitrogen losses, and a low-fiber diet that minimises metabolic fecal nitrogen loss. Because the bilby inhabits an environment where its food supply is unpredictable and of seasonally low nitrogen content, this species is likely to benefit from its low requirement for nitrogen.

Selective digesta retention and coprophagy in Brandt's vole (Microtus brandti).

Digestive performance, gut morphology and rate of digesta passage were measured in Brandt's voles (Microtus brandti) offered pelleted diets of low (25% neutral-detergent fibre) and high (38%) fibre content. Digestion coefficients of dry matter, crude fat, crude protein, energy and fibre were all significantly lower on the higher fibre diet. Although not significantly higher, dry matter intakes were more than maintained when extra cellulose was included in the diet, so that intakes of digestible energy were only 22% lower on the higher fibre diet. Total length and total gut tissue weight increased significantly, and the length and tissue weight of the caecum, proximal colon, and distal colon were significantly greater on the higher fibre diet as well. Total tract mean retention time (MRT) of a solute marker (Co-EDTA) was significantly greater than that of a particle marker (Cr-cell walls) on the lower fibre diet, and in the same direction on the higher fibre diet. The ratio of solute to particle MRTs (the solute/particle differential retention ratio) was 1.45 on the lower fibre diet and 1.19 on the higher fibre diet. There were no significant differences in marker MRTs between diets. Examination of marker concentrations in the stomach, small intestine, caecum and colon of voles killed at 0.5-h intervals after a pulse dose of Co-EDTA indicated that the marker was recycled to the stomach by coprophagy. Thus, as in other microtine rodents, an increase in gut capacity, selective digesta retention and recycling of digesta via coprophagy enables Brandt's voles to utilise diets of higher fibre content than may be expected for such a small (45 g) mammalian herbivore.

Effects of dietary fibre on digesta passage, nutrient digestibility, and gastrointestinal tract morphology in the granivorous Mongolian gerbil (Meriones unguiculatus).

To investigate digestive tract performance in Mongolian gerbils (Meriones unguiculatus), food intake and digestibility, digesta passage rate, and gastrointestinal tract morphology were measured in captive animals fed low- or high-fibre diets. We used two markers (Co-ethylene diamine tetra-acetic acid for solutes and Cr-mordanted cell walls for particles) to measure differential passage rates of digesta fractions in order to test for the presence of a colonic separation mechanism (CSM). Although dry-matter intakes on the high-fibre diet did not differ from those on the low-fibre diet, digestibilities of dry matter, neutral-detergent fibre, acid-detergent fibre, crude protein, and crude fat were all significantly lower on the high-fibre diet. Gross energy intake on the high-fibre diet also did not differ from that on the low-fibre diet, but energy lost in faeces was much higher than on the low-fibre diet; thus, energy digestibility and digestible energy intake were significantly lower on the high-fibre diet. The lengths and dry-tissue masses of all segments of the gastrointestinal tract tended to enlarge in response to increased dietary fibre, but only the total tract contents, contents of the small intestine, and length and dry-tissue mass of the caecum increased significantly. The mean retention time (MRT) of the particle marker was significantly greater than that of the solute marker on the low-fibre but not the high-fibre diet; the solute/particle differential retention ratio was 0.62 on the low-fibre diet and 0.90 on the high-fibre diet. Thus, there was no evidence for selective retention of the solute marker on either diet. The MRT of the particle marker was significantly lower on the high-fibre diet and in the same direction as the MRT of the solute marker. These results suggest that the granivorous Mongolian gerbil has no CSM but can adjust its digestive tract capacity to accommodate greater quantities of low-quality food.

The maintenance nitrogen requirement of the zebra finch Taeniopygia guttata.

A procedure is described for the accurate determination of the maintenance nitrogen requirement (MNR) of small granivorous birds. When used with the zebra finch (Taeniopygia guttata), it yielded a MNR of 403 mg kgW(-0.75) d(-1). This is lower than most other passerines so far measured and more similar to some nonpasserine species. Similarly, the value for endogenous nitrogen loss estimated for the zebra finch (153 mg kgW(-0.75) d(-1)) is less than that for passerines in general but higher than the nonpasserine value. We suggest that the low MNR of the strictly granivorous zebra finch is primarily an adaptation to seed diets in which high-quality protein is a limiting factor. Comparison with a wider range of species reported in the literature was restricted because inappropriate methods have been used to estimate MNR in many cases, including the use of growing or reproducing birds and the assumption that maintenance of body mass necessarily coincides with maintenance of zero nitrogen balance.

Digestive performance and digesta passage in the omnivorous greater bilby, Macrotis lagotis (Marsupialia: Peramelidae).

Omnivores such as the greater bilby (Macrotis lagotis) consume a variety of dietary items and often are faced with large changes in the nutrient composition of their food. This paper explores the basis for the dietary flexibility of the bilby by comparing digestive performance and digesta retention patterns of captive bilbies fed either an insect diet (mealworm larvae) or a plant diet (mixed seeds). Mean retention times (MRTs) of particle and solute markers in the gastrointestinal tract did not differ significantly between the two diets, but MRT of the particle marker was significantly longer than that of the solute marker on both the mealworm (particle: 23.5 h; solute: 17.9 h) and mixed seed (particle 33.0 h; solute: 30.2 h) diets. Lack of selective retention of solutes and small particles in the bilby gastrointestinal tract probably restricts them to relatively low-fibre diets, such as those based on seeds rather than leaves and stems of plants. It was observed radiographically that the major sites of digesta retention were the caecum, proximal colon and distal colon, and thus the hindgut is probably the principal site of microbial fermentation. The mealworms were more digestible than the mixed seeds, but digestible energy intake (mealworm: 939 kJ x kg(-0.75) x d(-1); mixed seed: 629 kJ x kg(-0.75) x d(-1)) was high enough for maintenance of body mass and positive nitrogen balance on both diets. Thus, although bilbies may be limited in their ability to utilize high-fibre diets by a lack of selective retention of solutes and small particles in their hindgut, their digestive strategy is flexible enough to accommodate at least some diets of both animal and plant origin. Such a strategy should benefit an animal inhabiting environments in which food resources are unpredictable in their relative abundance.

Responses of the digestive tract of the omnivorous northern brown bandicoot, Isoodon macrourus (Marsupialia: Peramelidae), to plant- and insect-containing diets.

The gastrointestinal tract of omnivores such as bandicoots (Marsupialia: Peramelidae) must be able to process foods as different as invertebrates, fungi and plant material. We studied the mechanisms involved in the utilisation by captive northern brown bandicoots (Isoodon macrourus) of insect larvae and milled lucerne (Medicago sativa) hay incorporated into a basal diet of a commercial small carnivore mix. Animals on the plant-basal mix digested less dry matter, energy, lipid, fibre and total nitrogen, but consumed 79% more dry matter than those on the insect-basal mix. Consequently intake of digestible energy (i.e. energy absorbed) was not significantly different between diets. Mean retention time (MRT, the mean time a marker remains in the tract) of a large particle marker was shorter on the plant-basal mix, reflecting its higher intake, but MRT of a solute marker was not significantly different between diets. Consequently the solute marker was retained longer than the particle marker on the plant-basal mix, indicating selective retention of solutes and very small particles in the caecum on this diet. This was confirmed by a higher proportion of small particles in the caecum than the distal colon of road-killed I. macrourus. Thus the main responses by I. macrourus to the plant-basal mix appeared to be an increase in gastrointestinal tract capacity (from radiographic evidence), selective retention of solutes and very small particles in the caecum, and facilitated passage of less tractable large particles through the colon. As a consequence, food intake was higher on the plant-basal mix, which compensated for its lower digestibility, and intake of digestible energy was similar to that on the insect-basal mix. This considerable flexibility of the morphologically rather simple digestive tract of northern brown bandicoots helps to explain their ability to cope with naturally variable diets consisting of mainly invertebrates in summer to much more plant and fungal material in winter, and to survive in nutritionally dynamic environments such as heathlands where there can be dramatic changes in food type and availability following periodic wildfires.

Contributions of microbes in vertebrate gastrointestinal tract to production and conservation of nutrients.

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.

Production of milk and nutrition of the dependent young of free-ranging koalas (Phascolarctos cinereus).

The production of milk by lactating females, and energy expenditure and foliage intake of their dependent young, were investigated in free-ranging koalas. Koalas had the lowest mass-specific daily milk-energy production at peak lactation so far recorded in a mammal, but the duration of reproduction was 58% longer than the combined marsupial and eutherian average. As a consequence, the total energy input to reproduction in koalas was similar to that in other mammals. We propose that the prolonged lactation and low daily rate of energy transfer to the young by female koalas is an adaptation to the low energy availability from their diet of Eucalyptus foliage. Energy requirements (field metabolic rates) of young koalas were lower than those expected for typical marsupials (only 60% at permanent pouch exit), which may be a necessary preadaptation that allows the low rate of maternal energy transfer. However, the energy requirements of the adult females were no lower than expected for marsupials. This pattern of energy requirements and age resulted in a linear relationship between field metabolic rate and mass for the koalas in this population. Differences in milk production between the years of the study coincided with fluctuations in the availability of preferred young foliage, which suggests that lactational output by koalas may be flexible and affected by diet quality. Despite the interannual differences in milk production, growth of the young was similar in the two years.

The diet and digestive energetics of an Australian short-necked turtle, Emydura macquarii.

We described the diet of Emydura macquarii, an omnivorous turtle from south-eastern Australia, compared its digestive performance on diets of fish or plants at two temperatures, and related how both diet and temperature affect its food selection in nature. Filamentous algae constituted 61% of the stomach content of E. macquarii. The turtles rarely fed on motile prey, but selected carrion from the lagoon bottom and terrestrial insects (Diptera, Hymenoptera and Coleoptera) trapped on the surface of the water. Digestive efficiency of E. macquarii was affected little by body temperature, in contrast to consumption rates and rates of passage which were strongly influenced by both temperature and diet. In combination, these responses resulted in a slower rate of digestion at 20 degrees C than at 30 degrees C. Digestive efficiency of E. macquarii on a herbivorous diet at 30 degrees C (49%) was about half that of turtles on a carnivorous diet (91%), but they had longer transit times (118 h on the plant diet versus 70 h). Lower consumption rates and longer mean retention times in turtles fed plants compared those fed fish relate to slower digestive processing of the plant. Rapid processing and higher consumption rates of fish by E. macquarii resulted in higher energy gains compared to turtles consuming plants (almost 100 times more energy at 30 degrees C). The laboratory results suggest that fish carrion and aquatic and terrestrial invertebrates are probably essential dietary items of E. macquarii in the wild, because its metabolic requirements cannot be met from aquatic macrophytes alone.

The digestive strategy of the common marmoset, Callithrix jacchus.

Digestive tract morphology and function were studied in the common marmoset (Callithrix jacchus), a small (350 g) exudivore with a well-developed caecum. Transit times (times of first appearance of the markers in the faeces following a pulse dose in the food) were similar for Co-EDTA, which marks the fluid phase of the digesta, and Cr-mordanted cell walls, which marked the large (600-1200 microns) particulate phase of the digesta. However, mean retention time (the average time taken for the markers to transverse the whole digestive tract) for Co-EDTA was significantly longer than for Cr-cell walls, indicating selective retention of fluid digesta relative to large particles, probably in the caecum. These data are consistent with a digestive strategy of the common marmoset that appears to be based on rapid digestion of higher quality foods (animal prey, fruits) in the small intestine, followed by microbial fermentation of the complex polysaccharides of plant exudates in the caecum, which would allow for considerable dietary flexibility in its natural habitat of scrub forests.

Selective retention of fluid digesta in the hindgut of bandicoots and other marsupial caecum fermenters.

Many small hindgut fermenters have a mechanism in the proximal colon that separates fluid and/or bacteria from large particles, and excretes the large particles relatively rapidly. The fluid and/or bacteria are retained in the caecum, which concentrates digestive effort in that region of the hindgut and improves overall digestive efficiency. Previously observed among marsupials only in arboreal marsupials, selective retention of fluid digesta has recently been reported also in bandicoots, small terrestrial omnivores. The separation mechanism operated independently of the nature of the diet, indicating that it is probably an important factor in the ability of bandicoots to switch between insect and plant foods, and thus to exploit nutritionally unpredictable environments. Results are discussed in relation to possible locations in the marsupial hindgut of the pacemaker that in eutherians has been shown to initiate retrograde movement of fluid and small particles in the proximal colon towards the caecum.

Digestive performance and selective digesta retention in the long-nosed bandicoot, Perameles nasuta, a small omnivorous marsupial.

Bandicoots are opportunistic omnivores that feed on invertebrates, fungi and both epigeal and hypogeal plant parts. We examined the performance of the digestive tract of the long-nosed bandicoot (Perameles nasuta) in terms of intake and total digestibility, patterns of excretion of inert digesta markers, and likely sites of digesta retention, on two diets designed to mimic part of their natural plant and insect diets. On the insect diet (mealworm larvae), bandicoots virtually maintained body mass at a digestible energy intake of 511 kJ.kg-0.75.day-1 and were in strongly positive nitrogen balance. In contrast, on the plant diet (shredded sweet potato), bandicoots ate only one-third as much digestible energy, lost 7% body mass, and were in negative nitrogen balance. Mean retention times of two particle markers on the plant diet (27.5 and 27.0 h) were more than double those on the insect diet (12.4 and 11.2 h), and on both diets the mean retention time of the fluid digesta marker was greater than those of the particle markers, indicating consistent selective retention of fluid digesta in the gut. It was seen radiographically than in mealworm-fed bandicoots major sites of digesta retention were the distal colon and rectum, whereas in the sweet potato-fed animals the caecum and proximal colon were principal sites.(ABSTRACT TRUNCATED AT 250 WORDS)

Acetate, butyrate and proline uptake in the caecum and colon of prairie voles (Microtus ochrogaster).

We have measured unidirectional uptake (not transmural flux) of acetate, butyrate and proline by everted sleeves of intact tissue from the jejunum, caecum, proximal colon and distal colon of prairie voles (Microtus ochrogaster). There was active (i.e. Na(+)-dependent) transport of L-proline in the jejunum, but we found no evidence for it in any region of the hindgut (i.e. the caecum, proximal colon and distal colon). Uptake of acetate was carrier-mediated in all three regions of the hindgut, but the Jmax and apparent Km (< or = 1.5 mmol l-1) were low, and uptake was primarily passive over the concentration range 10-50 mmol l-1, which spanned measured acetate levels in the caecum and proximal colon. At 100 mmol l-1, acetate uptake (nmol min-1 cm-2) was higher (P < 0.001) in distal colon (359 +/- 33) than in the proximal colon (225 +/- 17) and caecum (150 +/- 5) (mean +/- S.E., N = 8). Uptakes summed over the length of each region were also higher (P < 0.001) in the distal colon at 100 mmol l-1, but not at low concentrations (0.1 mmol l-1). Uptakes normalized to diffusion coefficients were higher for butyrate than acetate and were lowest for L-glucose (which is absorbed passively via an aqueous pathway) in all regions, indicating that uptake of the short-chain fatty acids involves solubilization in the lipid bilayer of the apical membrane. The short-chain fatty acids absorbed from the hindgut of the vole were equivalent to 22% of standard metabolic rate or 15% of resting (but fed) metabolic rate.

Effects of water restriction on digestive function in two macropodid marsupials from divergent habitats and the feral goat.

The effects of water restriction on digestive function in the euro (Macropus robustus erubescens) found in the arid zone of inland Australia, the eastern wallaroo (M.r.robustus) from more mesic regions of eastern Australia, and the feral goat (Capra hircus) found throughout the range of M. robustus, were compared in order to examine some physiological adaptations required by herbivores for the exploitation of arid environments. Eight animals of each species were held in individual metabolism cages in temperature-moderated rooms and given a chopped hay diet ad libitum. Half the animals were restricted to 40 ml water.kg-0.80.day-1. This was 40%, 32% and 57% of voluntary drinking water intake in the euro, wallaroo and goat, respectively. All species responded to water restriction by reducing faecal, urinary and evaporative water losses in association with reductions in feed intake. All animals increased urine osmolality and electrolyte concentrations but not to maximal levels, while packed-cell volume and plasma osmolality and electrolyte concentrations were unaffected by water restriction. The euro displayed a suite of characteristics that separated it from the wallaroo in terms of physiological adaptation, including lower voluntary water intake, an increase in fibre digestibility and maintenance of nitrogen balance during water restriction, and lower faecal water efflux associated with a consistently lower faecal water content (54% versus 59% water in the wallaroo during water restriction, P < 0.05). The euro's colon was 37% longer (P < 0.01) than that of the wallaroo. The goat had the lowest faecal water efflux (P < 0.05) and the longest colon (P < 0.001). Water restriction did not affect water content in digesta, nor short-chain fatty acid concentrations or production rates in vitro. Total body water, as a proportion of body mass, was depressed (P < 0.05) in the macropodids, but not in the goat. The reduction in dietary nitrogen intake, which accompanied water restriction, was partially compensated by an increase in urea degradation in the gut from 68% to 76% of urea synthesis water-restricted macropodids. These responses to water restriction are discussed in relation to those reported in other macropodid and ruminant species.

Digestive tract morphology and digestion in the wombats (Marsupialia: Vombatidae).

Wombats consume grasses and sedges which are often highly fibrous. The morphology of the digestive tract and the sequence of digestion were studied in two species of wombats from contrasting habitats: Vombatus ursinus from mesic habitats and Lasiorhinus latifrons from xeric regions. Studies were performed on wild wombats consuming their natural winter diets, and on captive wombats fed a high-fibre pelleted straw diet. Vombatus had a shorter digestive tract (9.2 vs 12.5 times body length) of greater capacity (wet contents 17.9 vs 13.7% body weight) than Lasiorhinus. The most capacious region of the digestive tract was the proximal colon (62-79% of contents). The proportional length and surface area of the proximal colon were greater in Vombatus, but those of the distal colon were greater in Lasiorhinus. These digestive morphologies may reflect adaptations for greater capacity and longer retention of digesta in Vombatus, but greater absorption and lower faecal water loss in Lasiorhinus. Apparent digestion along the digestive tract was estimated by reference to lignin. The proximal colon was the principal site of fibre and dry matter digestion, whereas nitrogen was mainly digested in the small intestine. Depot fats in captive wombats were highly unsaturated and reflected those in the diet. Therefore, lipids, proteins and soluble carbohydrates in the plant cell contents were digested and absorbed in the stomach and small intestine. Conversely, dietary fibre was probably retained and digested by microbial fermentation along the proximal colon.

Hindgut fermentation in the wombats: two marsupial grazers.

The wombats Vombatus ursinus and Lasiorhinus latifrons have a capacious proximal colon with only a vestigial caecum. The pattern of microbial fermentation in the hindgut of both species was studied in captive animals fed a pelleted straw diet and in wild wombats feeding on their natural winter diets. Digesta pH was low in the stomach but near neutrality along the hindgut, indicating effective absorption and/or buffering of the colonic contents. Initial proportions and production rates of short chain fatty acids in vitro reflected the fermentation of plant cell walls. Proportions of isobutyrate, isovalerate and n-valerate increased towards the distal colon indicating proteolysis and subsequent fermentation of amino acids. The low ammonia content of digesta fluid suggested that ammonia released from these amino acids was absorbed and utilized by the wombats and their gut microbes. Wild wombats had higher concentrations and production rates of short chain fatty acids than captive animals, which was consistent with the higher apparent digestibility of their natural diet. The energy from short chain fatty acids in captive animals was 30-33% of digestible intake. Energy intakes were low and similar to resting metabolic rates estimated for marsupials. Actual resting metabolic rates of the wombats are probably lower than these estimates, and the proportion of energy derived from fermentation substantially higher than the 53-61% estimated in wild wombats. The energy from fermentation clearly enables wombats to utilize diets high in fibre.

Digesta retention and fibre digestion in brushtail possums, ringtail possums and rabbits.

1. Mean retention times (MRTs) of fluid (marked with Co-EDTA), fine particles (mordanted with Yb) and large particles (mordanted with Cr) were measured in brushtail possums (Trichosurus vulpecula), ringtail possums (Pseudocheirus peregrinus) and laboratory rabbits fed semipurified diets. 2. In brushtail possums there were no significant differences in MRT among the three digesta markers. 3. In ringtail possums MRTs of the fluid and fine particle markers were approximately twice that of the large particle marker, indicative of selective retention of both fluid and fine particles in the caecum. 4. In the rabbit MRT of fine particles was also greater than that of large particles, again indicative of selective retention of fine particles in the caecum. 5. Fibre digestibility was greater in the rabbits than in the ringtail possums, and greater for neutral-detergent fibre (including agar) but less for acid-detergent fibre in the rabbits than in the brushtails. Differences in fibre digestibility between brushtails and rabbits were explained by differences in patterns of digesta flow. However, the higher digestibilities of fibre in the rabbits than in the ringtail possums could not be explained on a similar basis.

Energy expenditure and protein turnover in three species of wallabies (Marsupialia: Macropodidae).

Relationships between basal and fed metabolic rates and whole-body protein turnover rates were examined in three species of wallabies, the red-necked pademelon (Thylogale thetis), parma wallaby (Macropus parma) and tammar wallaby (M. eugenii). There were no significant differences among wallaby species in basal metabolic rate (BMR) which was 30% below eutherian mammals. However, the fed metabolic rate of the tammar was lower than that of the other two species (P less than 0.05), as was the protein turnover rate (P less than 0.01) which is consistent with its lower voluntary feed intake and with its lower maintenance nitrogen requirement. Protein turnover rates in the wallabies were 23-47% lower than in eutherian mammals. Similarly, protein synthesis made a lower contribution to fed metabolic rates in the wallabies (7-8%) than in eutherians (17-25%). Thus, compared with several eutherian species, macropodid marsupials have low rates of both energy and protein metabolism, but within the macropodids there is not necessarily a close link between basal metabolic rate and whole-body protein turnover.