SUMMARY Many large mammals show pronounced seasonal fluctuations of metabolic rate(MR). It has been argued, based on studies in ruminants, that this variation merely results from different levels of locomotor activity (LA), and heat increment of feeding (HI). However, a recent study in red deer (Cervus elaphus) identified a previously unknown mechanism in ungulates -nocturnal hypometabolism - that contributed significantly to reduced energy expenditure, mainly during late winter. The relative contribution of these different mechanisms to seasonal adjustments of MR is still unknown, however. Therefore, in the study presented here we quantified for the first time the independent contribution of thermoregulation, LA and HI to heart rate(fH) as a measure of MR in a free-roaming large ungulate,the Przewalski horse or Takhi (Equus ferus przewalskii Poljakow). f H varied periodically throughout the year with a twofold increase from a mean of 44 beats min-1 during December and January to a spring peak of 89 beats min-1 at the beginning of May. LA increased from 23% per day during December and January to a mean level of 53% per day during May, and declined again thereafter. Daily mean subcutaneous body temperature (Ts) declined continuously during winter and reached a nadir at the beginning of April (annual range was 5.8°C),well after the annual low of air temperature and LA. Lower Ts during winter contributed considerably to the reduction in fH. In addition to thermoregulation, fH was affected by reproduction, LA, HI and unexplained seasonal variation, presumably reflecting to some degree changes in organ mass. The observed phase relations of seasonal changes indicate that energy expenditure was not a consequence of energy uptake but is under endogenous control, preparing the organism well in advance of seasonal energetic demands.
Large ruminants respond to changing plant phenology during winter by decreasing voluntary food intake, increasing gut passage time and utilizing body fat reserves. It is uncertain, however, how other large mammals with a non-ruminant digestive physiology cope with winter forage conditions. Therefore, we investigated seasonality of energy intake in a large herbivorous wild mammal, the Przewalski horse (Equus ferus przewalskii). Throughout all seasons we used the n-alkane method to measure daily dry matter intake (DMI), diet composition and digestion, and determined an index of gut passage time in horses living under close to natural conditions. DMI correlated positively with its content of crude protein and nitrogen-free extract. Independent of these effects, DMI further varied seasonally with a peak in autumn and a nadir in late winter. Fluctuations of DMI corresponded to the annual change in body condition, which decreased during winter while energy reserves were depleted, and increased during the fattening period. Gut passage time varied in the course of the year and was longer during winter when the diet was high in crude fibre. Nevertheless, changes in gut passage time occurred rather independently of changes in forage composition and DMI, suggesting endogenous control for timely adaption of the digestive strategy to meet predictable changes in forage quality.
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