Scaling at different ontogenetic stages: Gastrointestinal tract contents of a marsupial foregut fermenter, the western grey kangaroo Macropus fuliginosus melanops.

Prominent ontogenetic changes of the gastrointestinal tract (GIT) should occur in mammals whose neonatal diet of milk differs from that of adults, and especially in herbivores (as vegetation is particularly distinct from milk), and even more so in foregut fermenters, whose forestomach only becomes functionally relevant with vegetation intake. Due to the protracted lactation in marsupials, ontogenetic differences can be particularly well investigated in this group. Here, we report body mass (BM) scaling relationships of wet GIT content mass in 28 in-pouch young (50 g to 3 kg) and 15 adult (16-70 kg) western grey kangaroos Macropus fuliginosus melanops. Apart from the small intestinal contents, in-pouch young and adults did not differ in the scaling exponents ('slope' in log-log plots) but did differ in the scaling factor ('intercept'), with an implied substantial increase in wet GIT content mass during the out-of-pouch juvenile period. In contrast to forestomach contents, caecum contents were elevated in juveniles still in the pouch, suggestive of fermentative digestion of milk and intestinal secretion residues, particularly in the caecum. The substantial increase in GIT contents (from less than 1 to 10-20% of BM) was associated mainly with the increase in forestomach contents (from 25 to 80% of total GIT contents) and a concomitant decrease in small intestine contents (from 50 to 8%), emphasizing the shifting relevance of auto-enzymatic and allo-enzymatic (microbial) digestion. There was a concomitant increase in the contents-to-tissue ratio of the fermentation chambers (forestomach and caecum), but this ratio generally did not change for the small intestine. Our study not only documents significant ontogenetic changes in digestive morpho-physiology, but also exemplifies the usefulness of intraspecific allometric analyses for quantifying these changes.

Ontogenetic scaling of the gastrointestinal tract of a marsupial foregut fermenter, the western grey kangaroo Macropus fuliginosus melanops.

As an animal grows, the relative sizes of their organs may grow proportionately or disproportionately, depending on ontogenetic changes in function. If organ growth is proportional (isometric), then the exponent of the scaling equation is 1.0. Relative decreases or increases in size result in exponents less than 1 (hypoallometric) or greater than 1 (hyperallometric). Thus, the empirical exponent can indicate potential changes in function. The entire gastrointestinal tract (GIT) of the foregut-fermenting western grey kangaroo Macropus fuliginosus melanops exhibited biphasic allometry across five orders of magnitude body mass (Mb; 52.0 g-70.5 kg). Prior to weaning at around 12 kg Mb, the entire empty GIT mass scaled with hyperallometry (Mb1.13), shifting to hypoallometry (Mb0.80) post-weaning. In addition, there were varying patterns of hyper-, hypo-, and isometric scaling for select GIT organs, with several displaying phase shifts associated with major life-history events, specifically around exit from the maternal pouch and around weaning. Mass of the kangaroo forestomach, the main fermentation site, scaled with hyperallometry (Mb1.16) before the stage of increased vegetation intake, and possibly after this stage (Mb1.12; P = 0.07), accompanied by a higher scaling factor (elevation of the curve) probably associated with more muscle for processing fibrous vegetation. The acid hindstomach mass showed hyperallometry (Mb1.15) before weaning, but hypoallometry (Mb0.58) post-weaning, presumably associated with decreasing intake of milk. Small intestine mass and length each scaled isometrically throughout ontogeny, with no discernible breakpoints at any life-history stage. The caecum and colon mass were steeply hyperallometric early in-pouch life (Mb1.59-1.66), when the young were ectothermic, hairless, and supported solely by milk. After around 295 g Mb, caecum mass remained hyperallometric (Mb1.14), possibly supporting its early development as a nidus for microbial populations to provide for secondary fermentation in this organ after the young transition from milk to vegetation.

An ecophysiologically informed model of seed dispersal by orangutans: linking animal movement with gut passage across time and space.

Fauna-mediated ecosystem service provision (e.g. seed dispersal) can be difficult to quantify and predict because it is underpinned by the shifting niches of multiple interacting organisms. Such interactions are especially complex in tropical ecosystems, including endangered peat forests of Central Borneo, a biodiversity hot spot and home to the critically endangered orangutan (Pongo pygmaeus wurmbii). We combined studies of the digestive physiology of captive orangutans in Australia with detailed field studies of wild orangutans in the Natural Laboratory of Peat-Swamp Forest of Sabangau, Central Kalimantan, Indonesia. By measuring the gut transit time (TT) of indigestible seed mimics (beads) in captivity and applying this as a temporal constraint to movement data of wild orangutans, we developed a mechanistic, time-explicit spatial model to project the seed dispersal patterns by these large-bodied, arboreal frugivores. We followed seven orangutans and established home range kernels using Time Local Convex Hull (T-LoCoH) modelling. This allowed us to model individual orangutan movements and to adjust these models according to gut transit times to estimate seed dispersal kernels. Female movements were conservative (core ranges of 55 and 52 ha in the wet and dry seasons, respectively) and revisitation rates to the same location of n = 4 in each 24-h block. Male movements were more unpredictable, yielding fragmented core ranges and revisitation rates to the same location of only 1.2 times each 24 h; males also demonstrated large disjunctions where they moved rapidly over long distances and were frequently lost from view. Seed dispersal kernels were nested predictably within the core ranges of females, but not males. We used the T-LoCoH approach to analyse movement ecology, which offered a powerful tool to predict the primary deposition of seeds by orangutans, thereby providing a reliable method for making a priori predictions of seed dispersal dynamics by other frugivores in novel ecosystems.

Effect of captivity on morphology: negligible changes in external morphology mask significant changes in internal morphology.

Captive breeding programmes are increasingly relied upon for threatened species management. Changes in morphology can occur in captivity, often with unknown consequences for reintroductions. Few studies have examined the morphological changes that occur in captive animals compared with wild animals. Further, the effect of multiple generations being maintained in captivity, and the potential effects of captivity on sexual dimorphism remain poorly understood. We compared external and internal morphology of captive and wild animals using house mouse (Mus musculus) as a model species. In addition, we looked at morphology across two captive generations, and compared morphology between sexes. We found no statistically significant differences in external morphology, but after one generation in captivity there was evidence for a shift in the internal morphology of captive-reared mice; captive-reared mice (two generations bred) had lighter combined kidney and spleen masses compared with wild-caught mice. Sexual dimorphism was maintained in captivity. Our findings demonstrate that captive breeding can alter internal morphology. Given that these morphological changes may impact organismal functioning and viability following release, further investigation is warranted. If the morphological change is shown to be maladaptive, these changes would have significant implications for captive-source populations that are used for reintroduction, including reduced survivorship.

Phenotypic plasticity in the common garden snail: big guts and heavier mucus glands compete in snails faced with the dual challenge of poor diet and coarse substrate.

Phenotypic plasticity allows animals to manage environmental challenges. Studies aimed at quantifying plasticity often focus on one challenge, such as diet, and one organ system, such the gastrointestinal tract, but this approach may not adequately reflect how plasticity could buffer multiple challenges. Thus, we investigated the outcomes of a dual challenge experiment that fed land snails either a high-fibre (low quality) or a low-fibre (high quality) diet, and simultaneously exercised them daily over 1.2 m on either a smooth surface of polyvinyl chloride (PVC) or a rough sandpaper. By the end of 20 days, snails fed the poor quality diet had a longer crop and oesophagus and a heavier intestine and rectum than those offered a low-fibre diet. Additionally, high-fibre fed snails had a smaller spermoviduct and oviduct. When also exercised on sandpaper, high-fibre fed snails had a smaller digestive gland, a main energy store, than those exercised on PVC. All snails exercised on sandpaper had a heavier pedal mucus gland, used a loping gait and used less mucus than those on PVC plastic, but there was no difference in the average speed of snails on either surface, supporting the conclusion that loping is a mucus conserving gait. Notably, snails faced with both a diet and substrate challenge had a smaller kidney, which could directly effect fecundity. This demonstrates that our dual challenge approach has potential for evaluating the costs and limits of the plasticity necessary to fully appreciate the evolutionary significance of plasticity in snails and other species.

Is "cooling then freezing" a humane way to kill amphibians and reptiles?

What is the most humane way to kill amphibians and small reptiles that are used in research? Historically, such animals were often killed by cooling followed by freezing, but this method was outlawed by ethics committees because of concerns that ice-crystals may form in peripheral tissues while the animal is still conscious, putatively causing intense pain. This argument relies on assumptions about the capacity of such animals to feel pain, the thermal thresholds for tissue freezing, the temperature-dependence of nerve-impulse transmission and brain activity, and the magnitude of thermal differentials within the bodies of rapidly-cooling animals. A review of published studies casts doubt on those assumptions, and our laboratory experiments on cane toads (Rhinella marina) show that brain activity declines smoothly during freezing, with no indication of pain perception. Thus, cooling followed by freezing can offer a humane method of killing cane toads, and may be widely applicable to other ectotherms (especially, small species that are rarely active at low body temperatures). More generally, many animal-ethics regulations have little empirical basis, and research on this topic is urgently required in order to reduce animal suffering.

Long-term changes in food availability mediate the effects of temperature on growth, development and survival in striped marsh frog larvae: implications for captive breeding programmes.

Food availability and temperature are known to trigger phenotypic change, but the interactive effects between these factors are only beginning to be considered. The aim of this study was to examine the independent and interactive effects of long-term stochastic food availability and water temperature on larval survivorship, growth and development of the striped marsh frog, Limnodynastes peronii. Larval L. peronii were reared in conditions of either constant or stochastic food availability and in water at three different temperatures (18, 22 and 26°C), and effects on survival, growth and development were quantified. Over the experimental period, larval growth rate was highest and survivorship lowest at the warmest temperature. However, changes in food availability mediated the effects of temperature, with slower larval growth and higher survivorship in stochastic food availability treatments. Tadpoles in the stochastic food availability treatments did not reach metamorphosis during the experimental period, suggesting that developmental stasis may have been induced by food restriction. Overall, these results demonstrate that changes in food availability alter the effects of water temperature on survival, growth and development. From an applied perspective, understanding how environmental factors interact to cause phenotypic change may assist with amphibian conservation by improving the number of tadpoles generated in captive breeding programmes.

Fibre-induced feed sorting in King Quail (Coturnix chinensis): behavioural plasticity elicited by a physiological challenge.

We examined the effect of an abrupt change in diet fibre content on the feed intake, gastrointestinal morphology and utilisation of gastroliths by a small (ca. 40 g body mass) herbivorous bird, the King Quail (Coturnix chinensis). King Quail were acclimated for 14 days on a low-fibre (LF) pullet starter diet. Following acclimation, half the quail population was immediately switched to a 23% wood-shaving diluted high-fibre (HF) diet for a further 14 days. Contrary to expectations, we found no differences in feed intake, gut morphology or gastrolith mass between the LF- and HF-fed quail. However, when switched from the LF to HF diet, the quail commenced feed-sorting behaviours that permitted HF-fed animals to maintain body condition (mass, abdominal fat mass) without adjustments to intestinal organ sizes or gastrolith mass. Feed sorting was initiated only after exposure to the HF diet, which corresponded with an immediate reduction in food intake, suggesting that the sorting behaviour was cued by a physiological challenge associated with the HF diet. This challenge apparently induced preferential sorting behaviour and was possibly due to abrupt changes in the rate of food passage, impacting satiation or other internal cues.

Is gastrointestinal plasticity in king quail (Coturnix chinensis) elicited by diet-fibre or diet-energy dilution?

Phenotypic plasticity of organ size allows some animals to manage fluctuations of resource quality or availability. Here, we examined the phenotypic plasticity of the gastrointestinal tract of king quail (Coturnix chinensis) in a diet-fibre manipulation study. Quail were offered either a control low-fibre (high-quality) food (8.5% neutral-detergent fibre; NDF), or one of two experimental diets of higher fibre contents of 16% NDF (i.e. low-quality food). To examine whether phenotypic plasticity of organ size was associated with the fibre content per se, or as a consequence of diluting the diet energy contents by adding fibre, one of the high-fibre feeds was 'balanced' with additional energy to match that of the low-fibre control diet. Total empty dry mass of the gastrointestinal tract was significantly heavier among birds offered the unbalanced high-fibre diet as compared with those offered the control diet, with birds offered the fibrous but energy-balanced diet having guts of intermediate size. The heavier entire-gut mass (dry) of quail offered the unbalanced high-fibre diet was associated mainly with these birds having significantly heavier gizzards. Notably, the larger gizzard in the birds offered the unbalanced high-fibre diet was associated with marked increases in their metabolisability (digestion) of diet fibre. Our findings suggest that the available energy in the diet may be more important for eliciting phenotypic changes in the gut of these herbivorous birds rather than simple physical effects of diet fibre on feed intakes or on muscular compensation to fibrous ingesta.

Physiology in conservation translocations.

Conservation translocations aim to restore species to their indigenous ranges, protect populations from threats and/or reinstate ecosystem functions. They are particularly important for the conservation and management of rare and threatened species. Despite tremendous efforts and advancement in recent years, animal conservation translocations generally have variable success, and the reasons for this are often uncertain. We suggest that when little is known about the physiology and wellbeing of individuals either before or after release, it will be difficult to determine their likelihood of survival, and this could limit advancements in the science of translocations for conservation. In this regard, we argue that physiology offers novel approaches that could substantially improve translocations and associated practices. As a discipline, it is apparent that physiology may be undervalued, perhaps because of the invasive nature of some physiological measurement techniques (e.g. sampling body fluids, surgical implantation). We examined 232 publications that dealt with translocations of terrestrial vertebrates and aquatic mammals and, defining 'success' as high or low, determined how many of these studies explicitly incorporated physiological aspects into their protocols and monitoring. From this review, it is apparent that physiological evaluation before and after animal releases could progress and improve translocation/reintroduction successes. We propose a suite of physiological measures, in addition to animal health indices, for assisting conservation translocations over the short term and also for longer term post-release monitoring. Perhaps most importantly, we argue that the incorporation of physiological assessments of animals at all stages of translocation can have important welfare implications by helping to reduce the total number of animals used. Physiological indicators can also help to refine conservation translocation methods. These approaches fall under a new paradigm that we term 'translocation physiology' and represent an important sub-discipline within conservation physiology generally.

Energy in-equivalence in Australian marsupials: evidence for disruption of the continent's mammal assemblage, or are rules meant to be broken?

The energy equivalence rule (EER) is a macroecological hypothesis that posits that total population energy use (PEU) should be independent of species body mass, because population densities and energy metabolisms scale with body mass in a directly inverse manner. However, evidence supporting the EER is equivocal, and the use of basal metabolic rate (BMR) in such studies has been questioned; ecologically-relevant indices like field metabolic rate (FMR) are probably more appropriate. In this regard, Australian marsupials present a novel test for the EER because, unlike eutherians, marsupial BMRs and FMRs scale differently with body mass. Based on either FMR or BMR, Australian marsupial PEU did not obey an EER, and scaled positively with body mass based on ordinary least squares (OLS) regressions. Importantly, the scaling of marsupial population density with body mass had a slope of -0.37, significantly shallower than the expected slope of -0.75, and not directly inverse of body-mass scaling exponents for BMR (0.72) or FMR (0.62). The findings suggest that the EER may not be a causal, universal rule, or that for reasons not yet clear, it is not operating for Australia's unique native fauna.

Caecal abnormality in a layer hen (Gallus gallus forma domestica) without deficits in digestive performance or egg productivity.

We report a case of a layer hen (Gallus gallus forma domestica) with deviation in the morphology of the caecum, and unique opportunity to investigate the digestive performance of the animal compared with normal hens. In a study investigating digestive and reproductive performance, an atypical caecal arrangement was found in a hen that was unremarkable in regards to body mass, digestive performance and egg productivity in comparison to other hens fed a similar diet. Examination of the gastrointestinal tract revealed a singular tubular outgrowth from the ileo-caecal junction, rather than the typical paired outgrowths. The single caecal duct bifurcated into two separate blind-ended sacs. Similar caecal deviations have been described in adult and juveniles, but no indications of animal performance were reported in these cases. We conclude that if the presence of an abnormal caecal arrangement reduces digestive abilities they were not obvious, and some compensatory mechanism/s may exist. Alternatively, the abnormal caecal arrangement of our hen might function adequately, such that no compensation in feed intakes or reduced egg productivity was required or observed.

Retention of different-sized particles and derived gut fill estimate in tammar wallabies (Macropus eugenii): physiological and methodological considerations.

The capacity of the digestive tract is an important parameter in understanding digestive adaptations, particularly in herbivores. Measures of this capacity ('gut fill') are commonly performed in killed animals, which has ethical and logistical implications. Alternatively, dry matter gut contents (DMC) can be estimated in live animals from food intake, digesta retention and digestibility, based on physical principles (Holleman and White, Can. J. Zool. 67, 488-494, 1989). Although this method has been used to some extent, it still awaits thorough validation. Here we estimated DMC in seven tammar wallabies during 5-day feeding trials and compared the results to those gained from dissections immediately after the trials. Calculated DMC exceeded that actually measured by 29 ± 22%. A closer inspection of the data suggested that this was partly due to the fact that DMC as measured by dissection is susceptible to short-term influences such as daily variation in food intake, whereas the calculated DMC represents an integrative measure over the whole period of the feeding trial. Correlations between both the measured digesta retention times, and the calculated DMC, with the measured wet contents mass suggest that it is particularly the DMC determined via dissection that needs to be measured with care. For a comparison of gut capacities, the calculated DMC therefore can be considered adequate, but should for a more widespread use be validated in further studies including more species and experimental regimes controlling food intake variation. Additionally, we tested whether very small (100-500 µm) and small (500-1000 µm) particles were retained differently in the tammar wallabies. There was no indication of such a difference. Whether the macropod forestomach selectively passes a certain particle fraction (that represents microbes) with the generally faster-passing fluids remains to be investigated with even smaller markers, e.g. labelled bacteria.

Coping with chaos: unpredictable food supplies intensify torpor use in an arid-zone marsupial, the fat-tailed dunnart (Sminthopsis crassicaudata).

The severity, duration and amplitude of extreme weather events are forecast to intensify with current climate trends, over both long (e.g. seasonal) and short (e.g. daily) time-scales. As such, the predictability of food supplies for many small endotherms is likely to become increasingly important. Numerous small mammals and birds combat food shortages using torpor, a controlled reduction in metabolic rate and body temperature that helps lower their daily energy requirements. As such, torpor often has been cited as a key feature allowing some small endotherms to survive highly unpredictable climates, such as tropics or dry deserts, but mensurative demonstrations of this are lacking. We have shown here that when a small desert marsupial, the fat-tailed dunnart (Sminthopsis crassicaudata), is offered unpredictable levels of daily food, they increase frequency of daily torpor and length of bouts compared with animals offered ad libitum food, but this was not found for animals offered a 75% [corrected] food-restricted diet. Our data suggest that simple food restriction may not be sufficient for evaluating the efficacy of torpor as a strategy for managing unpredictable climates.

Sensible heat loss from Muskoxen (Ovibos moschatus) feeding in winter: small calves are not at a thermal disadvantage compared with adult cows.

Muskoxen (Ovibos moschatus) are large (>200 kg adult body mass) mammalian herbivores that overwinter in the polar regions. Calves are around one-third the body mass of mature females and may be expected to suffer greater thermal stresses in winter compared with adults because the ratio of surface area to volume (SA:vol) is much greater for calves than for adults. We found that during feeding bouts, when animals are fully exposed to environmental conditions, calves did lose sensible (dry) heat more readily than adults (W m(-2)) in still air conditions. However, calves and cows lost less than 2%-6% of their estimated daily digestible energy intake as conductive, convective, and radiant heat losses accumulated during feeding bouts. More important, calves did not lose relatively more heat than larger adults in terms of sensible losses as part of their daily energy intake. Coat surface temperatures were only 2 degrees -5 degrees C above ambient even when air temperature fell to -40 degrees C. Body temperatures recorded deep within the ear canal near the tympanum fluctuated in both cows and calves. Muskoxen combine peripheral heterothermy and an exceptional winter coat to minimize sensible heat loss in winter. These mechanisms appear to have circumvented some of the thermal problems normally associated with a high SA:vol ratio in calves, which reflects the strong selection to conserve energy in winter.

Hindgut plasticity in wallabies fed hay either unchopped or ground and pelleted: fiber is not the only factor.

Phenotypic plasticity of the gastrointestinal tract is crucial for optimal food processing and nutrient balance in many vertebrate species. For mammalian herbivores, gut plasticity is typically correlated with the fiber content of forage; however, we show here that other factors such as ingesta particle size may effect profound phenotypic plasticity of the fermentative hindgut in a medium-sized (10-kg body mass) marsupial herbivore, the red-necked wallaby (Macropus rufogriseus). When dietary fiber contents were comparable, red-necked wallabies that were fed a finely ground, pelleted hay for 60-72 d had hindguts that were some 28% heavier (empty wet mass) than those fed unchopped hay. The hindguts of pellet-fed wallabies contained more wet ingesta, which was also of a finer particle size, than those fed hay, indicating some separation of large- and small-particle fermentation between the foregut and the hindgut, respectively. Such a digestive strategy would benefit animals by allowing fermentation of a range of ingesta particle sizes that are expected for free-ranging animals faced with a spectrum of diet types and qualities. The heavier hindgut of pellet-fed wallabies was correlated with increased concentrations of short-chain fatty acids (SCFAs) in the fermentative hindgut (cecum and proximal colon) and particularly with increases in the molar proportions of n-butyric acid. The mechanisms facilitating gut plasticity in herbivorous mammals are uncertain, but we suggest that manipulating ingesta particle size rather than dietary fiber could provide a useful tool for evaluating causal explanations. In particular, altering ingesta particle size could help to distinguish possible direct processes (e.g., the favoring of smaller intestinal microbes and production of specific SCFAs) from indirect affects of feed structure (e.g., muscular hypertrophy to compensate for increased intakes and digesta bulk or the fermentation of mucus secreted to promote the flow of viscous, fine-particle material).

Modelling digestive constraints in non-ruminant and ruminant foregut-fermenting mammals.

It has been suggested that large foregut-fermenting marsupial herbivores, the kangaroos and their relatives, may be less constrained by food intake limitations as compared with ruminants, due mainly to differences in their digestive morphology and management of ingesta particles through the gut. In particular, as the quality of forage declines with increasing contents of plant fibre (cellulose, hemicelluloses and lignin; measured as neutral-detergent fibre, NDF), the tubiform foregut of kangaroos may allow these animals to maintain food intakes more so than ruminants like sheep, which appear to be limited by fibrous bulk filling the foregut and truncating further ingestion. Using available data on dry matter intake (DMI, g kg(-0.75) d(-1)), ingesta mean retention time (MRT, h), and apparent digestibility, we modelled digestible dry matter intake (DDMI) and digestible energy intake (DEI) by ruminant sheep (Ovis aries) and by the largest marsupial herbivore, the red kangaroo (Macropus rufus). Sheep achieved higher MRTs on similar DMIs, and hence sheep achieved higher DDMIs for any given level of DMI as compared with kangaroos. Interestingly, MRT declined in response to increasing DMI in a similar pattern for both species, and the association between DMI and plant NDF contents did not support the hypothesis that kangaroos are less affected by increasing fibre relative to sheep. However, when DEI was modelled according to DDMIs and dietary energy contents, we show that the kangaroos could meet their daily maintenance energy requirements (MER) at lower levels of DMI and on diets with higher fibre contents compared with sheep, due largely to the kangaroos' lower absolute maintenance and basal energy metabolisms compared with eutherians. These results suggest that differences in the metabolic set-point of different species can have profound effects on their nutritional niche, even when their digestive constraints are similar, as was the case for these ruminant and non-ruminant foregut fermenters.

Could a big gut be too costly for muskoxen (Ovibos moschatus) in their first winter?

Young mammalian herbivores are more vulnerable to harsh winter conditions than adults, especially among large circumpolar species like the muskox (Ovibos moschatus). We compared feeding responses of muskox calves (body mass 95 kg) with those of mature, non-reproductive females (body mass 227 kg) in mid-winter when air temperatures fell to -40 degrees C. Food intakes (1.8-2.2 kg digestible dry matter (DM)d(-1)), digesta fill (27-32 kg wet mass) and digestibility of hay (52-58% of DM; 49-55% of gross energy) were similar between age groups even though calves were much smaller than adults. Calves fed more frequently (12 vs. 8 feeding bouts per day) and thus spent more time feeding each day than adults (387 vs. 343 min per day). High mass-specific food intakes of calves indicate higher requirements for maintenance of body tissue than adults, which could be related to a larger intestinal tract in young muskoxen. Notably, cows and calves maintained a constant body mass throughout, indicating that they were feeding at maintenance levels and that the relatively higher intakes of calves were not related to growth. Together, these data suggest that limited food availability due to snow cover or high animal density may reduce the survival of muskoxen in their first winter.

Ventilation patterns in red kangaroos (Macropus rufus Desmarest): juveniles work harder than adults at thermal extremes, but extract more oxygen per breath at thermoneutrality.

Juvenile mortalities in large mammals are usually associated with environmental extremes, but the basis for this vulnerability is often unclear. Because of their high surface area to volume ratio, juveniles are expected to suffer greater thermal stresses relative to adults. Coping with thermal stress requires the ventilatory system to accommodate increases in oxygen demand and respiratory water loss at thermal extremes. Because juveniles are smaller than adults, these demands may set up different constraints on their ventilatory system. Using red kangaroos (Macropus rufus Desmarest), an arid zone species, we compared the ventilatory capabilities of juveniles and adults at thermoneutral (25 degrees C) and extreme (-5 degrees C and 45 degrees C) ambient temperatures. We used an allometry to compare juvenile to adult ventilation, using predicted body mass scaling exponents for oxygen consumption (0.75), respiration rate (-0.25), tidal volume (1.0), ventilation rate (0.75) and oxygen extraction (0.0). At ambient 25 degrees C, the juveniles' resting metabolic rate was 1.6 times that of the mature females (ml min(-1) kg(-0.75)), accommodated by significantly higher levels of oxygen extraction of 21.4+/-1.8% versus 16.6+/-1.9% (P<0.05). At thermal extremes, juveniles showed typical mammalian responses in their ventilation, mirrored by that of adults, including higher metabolic and ventilation rates at ambient -5 degrees C and shallow panting at 45 degrees C. However, at thermal extremes the juvenile kangaroos needed to work harder than adults to maintain their body temperature, with higher rates of ventilation at ambient -5 degrees C and 45 degrees C, accomplished via larger breaths at -5 degrees C and higher respiratory rates at 45 degrees C.

Forage fibre digestion, rates of feed passage and gut fill in juvenile and adult red kangaroos Macropus rufus Desmarest: why body size matters.

Using red kangaroos Macropus rufus Desmarest, a large (>20 kg) marsupial herbivore, we compared the digestive capabilities of juveniles with those of mature, non-lactating females on high-quality forage (chopped lucerne Medicago sativa hay) of 43+/-1% neutral-detergent fibre (NDF) and poorer quality, high-fibre forage (chopped oaten Avena sativa hay) of 64+/-1% NDF. On chopped lucerne apparent dry matter (DM) digestibilities by young-at-foot (YAF) red kangaroos (an age that would normally be taking some milk from their mother), weaned juveniles and mature females were similar (55-59%). On chopped oaten hay apparent DM digestibility was lower in the YAF (35.9+/-2.3%) followed by weaned (43.4+/-2.8%) and mature females (44.6+/-1%). The digestion of NDF and its components (mainly cellulose and hemicellulose) was lowest among the YAF followed by weaned and then mature females. The YAF and weaned kangaroos could not sustain growth on the poor-quality diet, and appeared to be at or near maximal gut fill on both forages; the values being 114-122 g DM for YAF and 151-159 g DM for weaned kangaroos. Mean retention times (MRT) of particle and solute markers were significantly longer for the YAF and weaned kangaroos on oaten hay than on lucerne hay, and DM intake (g d(-1)) was approximately 50% lower on the oaten hay. In contrast, solute and particle MRTs in the mature females were not significantly affected by diet; they maintained DM intakes by increasing DM gut fill from 264+/-24 g on chopped lucerne to 427+/-26 g DM on chopped oaten hay. Clearly, the mature female kangaroos did not maximise gut fill on the high-quality forage, presumably as a consequence of their proportionally lower energy requirements compared with still-growing juveniles. Overall, we have provided the first mechanistic link between the physiological constraints faced by juvenile red kangaroos in relation to their drought-related mortalities, rainfall and forage quality.