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We evaluated the nutritional value of sporocarps of Rhizopogon vinicolor, a common hypogeous fungus in the coniferous forests of North America, for two small mammal species: the Californian red-backed vole (Clethrionomys californicus) and the northern flying squirrel (Glaucomys sabrinus). Although the nitrogen concentration of sporocarps was high, much of it was in non-protein form or associated with cell walls, suggesting that it may be of low nutritional value or protected from mammalian digestive enzymes. Sporocarps also had high concentrations of cell wall constituents, indicating low availability of digestible energy. When fed a diet of this fungus alone in a controlled feeding experiment both mammal species lost a small amount of body mass. Digestibilities of dry matter, nitrogen, cell wall constituents and energy from sporocarps by both species were lower than the digestibilities of other food types by other similarly sized small mammals. Red-backed voles digested the various components of sporocarps at least as well as the flying squirrels, even though they were almost six-fold smaller in body mass. This observation supports the notion that red-backed voles, like other microtine rodents, have morphological and physiological adaptations of the digestive system that are postulated to permit greater digestion of fibrous diets than predicted on the basis of body size. Despite this, our results re-affirm previous conclusions that hypogeous fungi are only of moderate nutritional value for most small, hindgut-fermenting mammals. Future studies should focus on the importance of mixed-species of fungi in the diet of small mammalian mycophagists.

Theory predicts that mammalian herbivores detoxify different classes of plant secondary compounds via separate metabolic pathways and that generalist herbivores maintain broad diet breadth to avoid overloading individual detoxification pathways. We tested the hypothesis that a generalist marsupial herbivore, the common brushtail possum, Trichosurus vulpecula, can maintain a higher intake of food when allowed to select from two diets containing different profiles of secondary compounds (phenolics and terpenes) than when given access to the diets individually. Diets consisted of a fruit and vegetable mash to which was added ground leaves of either Eucalyptus melliodora or E. radiata. E. melliodora and E. radiata differ in their concentrations and types of secondary compounds. Brushtail possums include these eucalypt species as part of their natural diet. We measured food consumption and detoxification metabolites of possums on these diets. Consistent with the hypothesis, animals presented with a choice of both diets consumed more food than animals given diets singly. One of the two indicators of detoxification, acid load in urine, differed significantly between diets while the other, glucuronic acid, did not. These results provide partial support for the hypothesis that diet breadth is governed by detoxification abilities.

Regulation of acid-base homeostasis is essential for mammals and birds. Biotransformation and metabolism of absorbed plant secondary metabolites (PSMs) results in the production of organic acids that threaten acid-base homeostasis. Consequently these acids must be buffered and excreted from the body. The production of an acid load from detoxified PSMs should occur in herbivorous mammals and birds and with most PSMs and so may provide a unifying theme to explain many effects of PSMs on animal metabolism. Since the organic acids will be largely ionized at physiological pH, disposal of the hydrogen ion and the organic anion may proceed independently. Most hydrogen ions (H(+)) from organic acids are eliminated by one or more of three ways: (1) when they react with bicarbonate in the extracellular fluid to form carbon dioxide and the carbon dioxide is exhaled, (2) when they bind to dibasic phosphate and are excreted by the kidney as monobasic phosphate, and (3) when they are buffered and retained in the skeletal system. The secretion of phosphate ions and ammonium excretion are two ways in which the kidney replaces bicarbonate ions that have been eliminated as carbon dioxide. Secretion in the kidney tubule is an important means of excreting excessive organic anions rapidly. This process is saturable and may be subject to competition from a variety of different metabolites. Lagomorphs have limited capacity to form new bicarbonate from ammonium excretion and may therefore be obliged to excrete other cations such as sodium to balance the excretion of organic anions from PSMs. Acidemia has wide-ranging impacts on animals but browsing mammals and birds may have to break down muscle tissues to provide for urinary ammonium in order to generate bicarbonate for buffering. Acidemia also can affect the extent of urea recycling. Animals consuming browse diets may have to regulate feeding so that the rate of formation of hydrogen ions does not exceed the rate of disposal. The mechanisms by which this could occur are unknown.

The metabolic pathways involved in degradation of tannin-protein complex (T-PC) were investigated in various facultatively anaerobic bacteria, with specific reference to fecal isolates from the koala including T-PC-degrading enterobacteria (T-PCDE),Streptococcus bovis, Klebsiella pneumoniae, andK. oxytoca. It was demonstrated that T-PCDE andS. bovis biotype I were capable of degrading protein complexed with gallotannin (a hydrolyzable tannin), but not that complexed with quebracho (a condensed tannin). Subsequent studies showed that these strains metabolized gallic acid to pyrogallol. Strains ofKlebsiella pneumoniae andK. oxytoca, which did not degrade T-PC, also metabolized gallic acid into pyrogallol. Pyrogallol was not degraded by any strains studied, but it was not detected in fresh feces of the koalas. The majority of strains isolated from feces could degrade phloroglucinol. Based on these findings, we propose that members of the gut microflora of the koala cooperate in the degradation of T-PC.

Optimal conditions for extraction of tannins and other phenolics from tree foliage and their subsequent storage rarely have been investigated. We investigated methods of drying leaves, optimal solvents, and the effects of light and temperature on the extractability and stability of condensed tannins (proanthocyanidins) and total phenolics from leaves ofEucalyptus trees. Aqueous acetone was a better solvent than aqueous methanol for condensed tannins and total phenolics, but condensed tannins were less stable in aqueous acetone than aqueous methanol. Stability of condensed tannins also was decreased substantially by room temperature versus 4°C and by exposure to indirect sunlight, although the assay for total phenolics was unaffected. For quantitative estimation of condensed tannins, extraction with 50% acetone was better than methods of direct analysis of leaf tissue. The highest estimates of total condensed tannins were obtained by exhaustive extraction with 50% acetone followed by direct analysis of the residue. Lyophilization of fresh leaf increased yield of condensed tannin (although usually by less than 10%). Lyophilization and subsequent storage of extracts had little effect on assays for condensed tannins or total phenolics.

The passage of particulate and solute digesta through the alimentary tract of koalas fed the mature foliage ofEucalyptus punctata was investigated using radioisotopic markers. The solute marker was retained for longer than the particulate marker, the mean retention times of both markers (213 h and 99 h respectively) being longer than has been reported in most other mammals. Both the pattern of excretion in faeces of orally administered markers and data from a slaughter experiment indicated that the principal site of marker retention was the highly developed caecum/proximal colon which appeared to function as a single mixing compartment. An undettermined physiological mechanism caused solutes and fine particles of digesta to be selectively retained in the caecum/proximal colon. It is suggested that this caused the particulate marker to distribute in two pools situated in parallel producing a marker excretion curve different from any reported in other mammals. It is postulated that the rate of clearance of dietary fibre from the hindgut would be maximized by this mechanism and that this in turn would elevate the upper limit to intake of food, and hence digestible nutrient, by koalas. Such an adaptation would partially compensate for the physiological constraints on nutrient acquisition from fibrous diets imposed by small body size and thus may be important in enabling the koala to utilize a diet avoided by most other small herbivores.

Digestion and energy metabolism in an arboreal marsupial, the koalaPhascolarctos cinereus, fed mature foliage from a common food tree, the grey gumEucalyptus punctata, were investigated. Six feeding (balance) experiments, at various times of year, and one slaughter experiment were performed and average daily oxygen consumption was measured. The average apparent digestibilities of dietary constituents were: dry matter 54%; total cell-contents 69%; available carbohydrate 92%; crude lipid 43%; total nitrogen 45%; total phenolics 91%; total cell walls 25%; hemicellulose 24%; acid-detergent fibre 25%; cellulose 31%; lignin 19%. Average digestible and metabolizable energy intakes were 0.50 and 0.43 MJ kg−0.75 d−1 respectively of which only 0.28 MJ kg−0.75 d−1 was expended in oxidative metabolism. The digestible energy intake required for maintenance was estimated to be 0.33 MJ kg−0.75 d−1, which is lower than that of eutherian and of other marsupial herbivores. The principal sources of metabolizable energy were non-structural carbohydrate and lipid. It is postulated that the ability of koalas to utilizeEucalyptus foliage as a sole source of nutrients is facilitated by their low requirement for energy and their ability to maximize intake of non cell-wall constituents.E. punctata foliage has a high digestible energy content compared with the foliage of many other trees and this may be a factor in its selection by koalas.

The rate of volatile fatty acid (VFA) production in the hindgut of three koalas (Phascolarctos cinereus) maintained onEucalyptus punctata foliage was measured by in vitro incubation.