Seasonal energy intake of Tibetan sheep on the harsh Qinghai-Tibetan Plateau (QTP) fluctuates greatly and is often well below maintenance requirements. The aim of this study was to gain insight into how the hypothalamus regulates energy homoeostasis in Tibetan sheep. We compared Tibetan and Small-tailed Han sheep (n 24 of each breed), which were each allocated randomly into four groups and offered one of four diets that differed in digestible energy densities: 8·21, 9·33, 10·45 and 11·57 MJ/kg DM. Sheep were weighed every 2 weeks, and it was assumed that the change in body weight (BW) reflected the change in energy balance. The arcuate nucleus of the hypothalamus in Tibetan sheep had greater protein expressions of neuropeptide Y (NPY) and agouti-related peptide (AgRP) when in negative energy balance, but lesser protein expressions of proopiomelanocortin (POMC) and cocaine and amphetamine-regulated transcript (CART) when in positive energy balance than Small-tailed Han sheep. As a result, Tibetan sheep had a lesser BW loss when in negative energy balance and stored more energy and gained more BW when in positive energy balance than Small-tailed Han sheep with the same dietary intake. Moreover, in the hypothalamic adenosine monophosphate-activated protein kinase (AMPK) regulation pathway, Tibetan sheep had greater adenosine monophosphate-activated protein kinase-α 2 protein expression than Small-tailed Han sheep, which supported the premise of a better ability to regulate energy homoeostasis and better growth performance. These differences in the hypothalamic NPY/AgRP, POMC/CART and AMPK pathways between breeds conferred an advantage to the Tibetan over Small-tailed Han sheep to cope with low energy intake on the harsh QTP.

Sheep are grazers and goats are intermediate feeders. By employing O2 consumption and heart rate measurements, resting metabolic rate (RMR) and field metabolic rate (FMR) were determined in four male fat-tailed Awassi sheep (44.0 ± 3.94) and four male Baladi goats (35.5 ± 5.42 kg) that were co-grazing natural pasture in the Negev Desert. There were 67.7 ± 3.75 g DM/m2 of herbaceous vegetation biomass, which was rapidly becoming senescent and more fibrous. We hypothesized that FMR of these desert-adapted ruminants would be relatively low when compared to other sheep and goat breeds, as animals in arid areas tend to have low metabolic rates. Both sheep (n = 6) and goats (n = 6) foraged 71% of the allotted 11 h free-pasture period; however, sheep grazed more than goats (P < 0.001); whereas goats browsed more than sheep (P < 0.001). RMR was higher (P = 0.007) in sheep than in goats (529 ± 23.5 v. 474 ± 25.4 kJ/kg0.75 BW/d), but FMR did not differ between species (618 ± 55.7 v. 613 ± 115.2 kJ/kg0.75 BW/d). In addition, the cost of activities, as a proportion of FMR, did not differ between sheep and goats; FMR increased by 89 kJ/kg0.75 BW/d or 17% in sheep and by 138 kJ/kg0.75 BW/d or 29% in goats. In comparing FMRs of sheep and goats in this study with these species in other studies, differences were inconsistent and, therefore, our hypothesis was not supported.

This study aimed to gain insight into how adipose tissue of Tibetan sheep regulates energy homoeostasis to cope with low energy intake under the harsh environment of the Qinghai-Tibetan Plateau (QTP). We compared Tibetan and Small-tailed Han sheep (n 24 of each breed), all wethers and 1·5 years of age, which were each divided randomly into four groups and offered diets of different digestible energy (DE) densities: 8·21, 9·33, 10·45 and 11·57 MJ DE/kg DM. When the sheep lost body mass and were assumed to be in negative energy balance: (1) adipocyte diameter in subcutaneous adipose tissue was smaller and decreased to a greater extent in Tibetan than in Small-tailed Han sheep, but the opposite occurred in the visceral adipose tissue; (2) Tibetan sheep showed higher insulin receptor mRNA expression and lower concentrations of catabolic hormones than Small-tailed Han sheep and (3) Tibetan sheep had lower capacity for glucose and fatty acid uptake than Small-tailed Han sheep. Moreover, Tibetan sheep had lower AMPKα mRNA expression but higher mammalian target of rapamycin mRNA expression in the adipocytes than Small-tailed Han sheep. We concluded that Tibetan sheep had lower catabolism but higher anabolism in adipose tissue and reduced the capacity for glucose and fatty acid uptake to a greater extent than Small-tailed Han sheep to maintain energy homoeostasis when in negative energy balance. These responses provide Tibetan sheep with a high ability to cope with low energy intake and with the harsh environment of the QTP.

The nutritional intake of Tibetan sheep on the harsh Qinghai–Tibetan Plateau is often under maintenance requirements, especially during the long, cold winter. However, they have adapted well and even thrive under these conditions. The aim of the present study was to gain insight into how the rumen epithelium of Tibetan sheep has adapted to the consumption of low-energy-level diets. For this purpose, we compared Tibetan and small-tailed Han sheep (n 24 of each breed, all wethers and 1·5 years of age), which were divided randomly into one of four groups and offered ad libitum diets of different digestible energy (DE) densities: 8·21, 9·33, 10·45 and 11·57 MJ DE/kg DM. The Tibetan sheep had higher rumen concentrations of total SCFA, acetate, butyrate and iso-acids but lower concentrations of propionate than small-tailed Han sheep. The Tibetan sheep had higher absorption capability of SCFA due to the greater absorption surface area and higher mRNA expression of the SCFA absorption relative genes than small-tailed Han sheep. For the metabolism of SCFA in the rumen epithelium, the small-tailed Han sheep showed higher utilisation of the ketogenesis pathway than Tibetan sheep; however, Tibetan sheep had greater regulation capacity in SCFA metabolism pathways. These differences between breeds allowed the Tibetan sheep to have greater capability of absorbing SCFA and better capacity to regulate the metabolism of SCFA, which would allow them to cope with low energy intake better than small-tailed Han sheep.

We investigated offspring quality in fleas (Xenopsylla ramesis) feeding on non-reproducing, pregnant or lactating rodents (Meriones crassus) and asked whether (a) quality of flea offspring differs dependent on host reproductive status; (b) fleas trade off offspring quantity for quality; and (c) quality variables are inter-correlated. Emergence success was highest when parents exploited pregnant hosts, while development time was longest when parents exploited lactating hosts. Male offspring from fleas fed on non-reproductive and pregnant hosts were larger than those from lactating hosts whereas female offspring from fleas fed on pregnant hosts were larger than those from both lactating and non-reproductive hosts. Male offspring survived under starvation the longest when their parents exploited lactating hosts and the shortest when their parents exploited pregnant hosts. Female offspring of parents that exploited lactating hosts survived under starvation longer than those that exploited non-reproductive and pregnant hosts. Emergence success and development time decreased as mean number of eggs laid by mothers increased. Fleas that were larger and took longer to develop lived significantly longer under starvation. These results indicate the presence of a trade-off between offspring quantity and quality in fleas exploiting female Sundevall's jird in varying reproductive condition but this trade-off depended on the quality trait considered.

We investigated spatial variation in the phylogenetic structure of host spectra in fleas parasitic on small mammals. Measures of phylogenetic host specificity ((phylogenetic species clustering (PSC) and phylogenetic species variability (PSV)) varied significantly more between than within flea species, but the proportion of variation which accounted for among-species differences was low. In 13 of 18 common flea species, at least one of the indices of the phylogenetic structure of regional host spectra revealed a significantly positive association with the phylogenetic structure of regional host assemblage, while relationships between PSC or PSV of the regional host spectrum and the distance from either the region of a flea's maximal abundance or latitude were not supported. Overall, results of this study demonstrated that although the degree of phylogenetic host specificity in fleas can be considered as a true attribute of a flea species, it is highly spatially variable, with phylogenetic structure of the surrounding host pool being the main reason behind this variation.

We studied the energy cost of egg production in two flea species (Parapulex chephrenis and Xenopsylla ramesis) feeding on principal (Acomys cahirinus and Meriones crassus, respectively) and auxiliary (M. crassus and A. cahirinus, respectively) rodent hosts. We predicted that fleas feeding on principal as compared with auxiliary hosts will (a) expend less energy for egg production; (b) produce larger eggs and (c) live longer after oviposition. Both fleas produced more eggs and spent less energy per egg when exploiting principal hosts. Parapulex chephrenis produced larger eggs after exploiting auxiliary hosts, while the opposite was true for X. ramesis. After oviposition, P. chephrenis fed on the auxiliary hosts survived for a shorter time than those fed on the principal hosts, while in X. ramesis the survival time did not differ among hosts. Our results suggested that one of the proximate causes for lower reproductive performance and subsequent lower abundance of fleas on auxiliary hosts is the higher energy cost of egg production. However, in some species, lower offspring number may be compensated to some extent by their size, although this compensation may also compromise their future reproduction via decreased survival. In addition, the reproductive strategy of exploitation of low profitable (i.e. auxiliary) hosts may differ between flea species.

We tested for the effect of age of a rodent host (Meriones crassus) on reproductive performance of fleas in terms of number and quality of offspring and predicted that fleas would perform better on juvenile and old than on subadult and adult hosts. The number of flea offspring was evaluated via egg and new imago production, while their quality was estimated via duration of development, resistance to starvation and body size. Although fleas produced more eggs when they exploited adults than when they exploited juvenile, subadult and old hosts, significantly more new imago emerged from fleas fed on juvenile and old hosts than on subadult and adult hosts. Fleas performed better when they fed on juvenile and/or old hosts than on subadult and adult hosts in 2 of 3 measures of offspring quality (duration of development and body size). Nevertheless, when offspring quality was estimated via resistance to starvation of a new imago, fleas demonstrated good performance in young (juvenile and subadult) hosts, while they performed poorly in old hosts. Thus, general reproductive performance of fleas was better when they exploited young and old hosts than when they exploited median age cohorts. However, the effect of host age on flea reproductive performance was manifested somewhat differently between (a) male and female hosts and (b) male and female flea offspring.

We asked if and how feeding performance of fleas on an auxiliary host is affected by the phylogenetic distance between this host and the principal host of a flea. We investigated the feeding of 2 flea species, Parapulex chephrenis and Xenopsylla ramesis, on a principal (Acomys cahirinus and Meriones crassus, respectively) and 8 auxiliary host species. We predicted that fleas would perform better (higher proportion of fleas would feed and take larger bloodmeals) on (a) a principal rather than an auxiliary host and (b) auxiliary hosts phylogenetically closer to a principal host. Although feeding performance of fleas differed among different hosts, we found that: (1) fleas did not always perform better on a principal host than on an auxiliary host; and (2) flea performance on an auxiliary host was not negatively correlated with phylogenetic distance of this host from the principal host. In some cases, fleas fed better on hosts that were phylogenetically distant from their principal host. We concluded that variation in flea feeding performance among host species results from interplay between (a) inherent species-specific host defence abilities, (b) inherent species-specific flea abilities to withstand host defences and (c) evolutionary tightness of association between a particular host species and a particular flea species.

We asked whether the age of a rodent host affects the feeding performance of fleas. We predicted that fleas would perform better on young and old than on adult rodents. To test this prediction, we determined bloodmeal size, rate of digestion and time of survival after a single bloodmeal in Xenopsylla ramesis feeding on Meriones crassus of different ages. Fleas took less blood from subadult and adult than from juvenile and old animals. Fleas digested blood of old hosts at the highest rate and blood of juvenile hosts most slowly. After a bloodmeal, fleas survived the longest if fed on a juvenile host. The effect of host gender on bloodmeal size and survival after a bloodmeal was manifested only in (a) subadult and adult hosts and (b) subadult hosts, respectively. Host gender affected the rate of digestion differently among digestion stages and host age cohorts. We explain the observed patterns in flea performance in terms of changes in the host's immune defences and nutritional quality of blood during the host's individual life. Our observations suggested that a host of a particular age could not be unequivocally predicted to be more or less beneficial for a parasite than a younger or an older host.

We studied the effect of density of larvae on pre-imaginal development in 2 flea species (Xenopsylla conformis and Xenopsylla ramesis) parasitic on 2 desert rodent species (Dipodillus dasyurus, adult body mass 20 g and Meriones crassus, 80 g). We predicted a decrease in duration of development with an increase in density of larvae. In general, in both flea species, duration of larva-to-pupa development decreased with an increasing larval density. In addition, this stage of development was longer in male fleas and in fleas from parents fed on D. dasyurus. The effect of larval density on larval development was manifested mainly when parent fleas fed on D. dasyurus. Duration of pupation decreased with increasing larval density only in offspring of fleas fed on G. dasyurus. In both fleas, pupation was longer in males. The effect of parent host on duration of pupation was found in X. ramesis only (longer if the host was M. crassus). Resistance of newly emerged fleas to starvation depended mainly on parent host species. Young X. conformis survived longer if their parents fed on D. dasyurus, whereas young X. ramesis survived longer if their parents fed on M. crassus. It was also found that (a) an individual flea that spent more time as a larva also spent more time as a pupa and (b) longer larval development resulted in a shorter time that a newly emerged flea was able to survive when starved.

Large areas of the Central Asian Republic of Kazakhstan are classified as desert and semi-desert rangelands and are subjected to severe climatic extremes. The sparse vegetation of these rangelands supports mainly grazing sheep, which are raised for wool and meat production. These two products are important to the national economy and the livelihood of the rural population. The extremes of climate cause periodic shortages of fodder availability that, together with the lack of drinking water and adequate animal shelters, can cause severe animal loss, particularly during the winter but also in other seasons. Desert rangeland improvements have been instigated by the introduction and planting out of palatable drought-resistant fodder shrubs to augment the natural species, many of which appear to have survived only because they are unpalatable and/or inedible. To determine the edibility of 15 natural species, Karakul sheep were observed grazing and/or browsing them and identified those species that were edible. It was predicted that the sheep would selectively browse from shrubs with low phenolic concentration, high estimated metabolizable energy content, high crude protein content and low fibre content. To test this prediction, the shrub foliage was analysed and edibility by sheep was related to chemical composition and estimated metabolizable energy content. Six of the 15 plant species were edible. The only significant factor in determining whether plants were eaten was total phenolic content – plants eaten had a content of 17·7±6·3 g/kg DM whereas plants not eaten had a content of 66·4±129·6 g/kg DM. Metabolizable energy, protein and fibre contents did not affect edibility of the plants.

Body mass was measured and body composition and energy requirements were estimated in sheep at four air temperatures (0 °C to 30 °C) and at four levels of energy offered (4715 to 11785 kJ/day) at a time when the sheep reached a constant body mass. Final body mass was affected mainly by metabolizable energy intake and, to a lesser extent, by air temperature, whereas maintenance requirements were affected mainly by air temperature. Mean energy requirements were similar and lowest at 20 °C and 30 °C (407·5 and 410·5 kJ/kg0·75, respectively) and increased with a decrease in air temperature (528·8 kJ/kg0·75 at 10 °C and 713·3 kJ/kg0·75 at 0 °C). Absolute total body water volume was related positively to metabolizable energy intake and to air temperature. Absolute fat, protein and ash contents were all affected positively by metabolizable energy intake and tended to be related positively to air temperature. In proportion to body mass, total body water volume decreased with an increase in metabolizable energy intake and with an increase in air temperature. Proportionate fat content increased with an increase in metabolizable energy intake and tended to increase with an increase in air temperature. In contrast, proportionate protein content decreased with an increase in metabolizable energy intake and tended to decrease with an increase in air temperature. In all cases, the multiple linear regression using both air temperature and metabolizable energy intake improved the fit over the simple linear regressions of either air temperature or metabolizable energy intake and lowered the standard error of the estimate. The fit was further improved and the standard error of the estimate was further lowered using a polynomial model with both independent variables to fit the data, since there was little change in the measurements between 20 °C and 30 °C, as both air temperatures were most likely within the thermal neutral zone of the sheep. It was concluded that total body energy content, total body water volume, fat and protein content of sheep of the same body mass differed or tended to differ when kept at different air temperatures.