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Summary 0[ Theoretical studies of large herbivore foraging assume that total daily grazing time is a key constraint on daily intake and diet choice[ We experimentally tested this assumption and investigated the e}ects of food availability on the ability of grazing sheep to compensate for restriction of available daily grazing time[ 1[ Foraging behaviour\ intake and diet digestibility by sheep\ were measured on grass pastures in a replicated 1 × 1 factorial experiment\ in which overnight access to pasture was varied "restricted overnight and continuous access# on two sward heights "4=4 and 2=9 cm#\ representing high and low food availability[ 2[ Regardless of food availability\ the overnight!restricted sheep fed for almost all of the available grazing time by grazing for fewer\ longer foraging bouts\ but still had much shorter total daily grazing time than the continuous access sheep[ 3[ In response to overnight penning\ the sheep had a signi_cantly higher instantaneous rate of intake achieved mainly via larger bites[ The continuous access sheep were hence not maximizing their short!term rate of intake\ whilst grazing according to the daily schedule considered normal for sheep[ 4[ The behavioural responses to overnight food restriction were able to counteract the reduction in daily grazing time only where food availability was high[ In contrast on short swards overnight grazing restriction led to a reduction in total daily intake[ We suggest that the interactions between the factors considered as constraints on foraging behaviour of herbivores are\ as yet\ only poorly quanti_ed[ Key!words] foraging constraint\ intake rate\ sheep\ sward height\ time limitation[ Journal of Animal Ecology "0888# 57\ 76Ð82

BackgroundGene duplication is a key evolutionary mechanism providing material for the generation of genes with new or modified functions. The fate of duplicated gene copies has been amply discussed and several models have been put forward to account for duplicate conservation. The specialization model considers that duplication of a bifunctional ancestral gene could result in the preservation of both copies through subfunctionalization, resulting in the distribution of the two ancestral functions between the gene duplicates. Here we investigate whether the presumed bifunctional character displayed by the single branched chain amino acid aminotransferase present in K. lactis has been distributed in the two paralogous genes present in S. cerevisiae, and whether this conservation has impacted S. cerevisiae metabolism.Principal FindingsOur results show that the KlBat1 orthologous BCAT is a bifunctional enzyme, which participates in the biosynthesis and catabolism of branched chain aminoacids (BCAAs). This dual role has been distributed in S. cerevisiae Bat1 and Bat2 paralogous proteins, supporting the specialization model posed to explain the evolution of gene duplications. BAT1 is highly expressed under biosynthetic conditions, while BAT2 expression is highest under catabolic conditions. Bat1 and Bat2 differential relocalization has favored their physiological function, since biosynthetic precursors are generated in the mitochondria (Bat1), while catabolic substrates are accumulated in the cytosol (Bat2). Under respiratory conditions, in the presence of ammonium and BCAAs the bat1Δ bat2Δ double mutant shows impaired growth, indicating that Bat1 and Bat2 could play redundant roles. In K. lactis wild type growth is independent of BCAA degradation, since a Klbat1Δ mutant grows under this condition.ConclusionsOur study shows that BAT1 and BAT2 differential expression and subcellular relocalization has resulted in the distribution of the biosynthetic and catabolic roles of the ancestral BCAT in two isozymes improving BCAAs metabolism and constituting an adaptation to facultative metabolism.

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The effects of four diets differing in their for age: concentrate ratio (80:20, 60:40, 40: 60 and 20:80; g/100 g fresh matter) on rumen characteristics, digestibility and digesta flow kinetics were investigated. Alfalfa hay was used as forage and concentrate was composed of barley, soya-bean meal and maize. Diets were prepared by mixing all ingredients and offered to the animals as complete diets. Eight mature Merino sheep, each fitted with a rumen cannula, were offered 1·055 kg dry matter per day of the corresponding diet over two experimental periods. The daily evolution of ruminai pH, volatile fatty acids (VFA) and ammonia nitrogen (N) concentrations were measured. Digestibility was determined by total faecal collection and Cr and Co were used as markers to estimate digesta passage rates. Microbial nitrogen flow at the duodenum (MNDF) was estimated from the urinary excretion of purine derivatives (PD). The apparent digestibility of organic matter increased (P < 0·001) whereas that of all fibrous fractions decreased linearly (P < 0·05) as the proportion of concentrate in the diet increased. Rumen pH decreased linearly (P < 0·001) with increasing proportions of concentrate but total VFA concentrations were unaffected by changes in the diet (P > 0·05). Both liquid and solid digesta outflow rates from the rumen decreased quadratically (P < 0·01) as the proportion of concentrate in the diet increased. The urinary excretion of total N, urea-N and ammonia-N was unaffected (P > 0·05) by changes in the diet. In contrast, the daily urinary excretion of both allantoin and total PD increased quadratically (P < 0·05) with increasing proportions of concentrate. Consequently, the estimated MNDF increased linearly (P < 0·001) from 9·9 g/day on the high forage diet to 14·5 g/day on the high concentrate diet.