Monitoring the prevalence and abundance of parasites over time is important for addressing their potential impact on host life histories, immunological profiles and their influence as a selective force. Only long-term ecological studies have the potential to shed light on both the temporal trends in infection prevalence and abundance and the drivers of such trends, because of their ability to dissect drivers that may be confounded over shorter time scales. Despite this, only a relatively small number of such studies exist. Here, we analysed changes in the prevalence and abundance of gastrointestinal parasites in the wild Soay sheep population of St. Kilda across 31 years. The host population density (PD) has increased across the study, and PD is known to increase parasite transmission, but we found that PD and year explained temporal variation in parasite prevalence and abundance independently. Prevalence of both strongyle nematodes and coccidian microparasites increased during the study, and this effect varied between lambs, yearlings and adults. Meanwhile, abundance of strongyles was more strongly linked to host PD than to temporal (yearly) dynamics, while abundance of coccidia showed a strong temporal trend without any influence of PD. Strikingly, coccidian abundance increased 3-fold across the course of the study in lambs, while increases in yearlings and adults were negligible. Our decades-long, intensive, individual-based study will enable the role of environmental change and selection pressures in driving these dynamics to be determined, potentially providing unparalleled insight into the drivers of temporal variation in parasite dynamics in the wild.

Research on the causes of sheep death in sea voyages from Australia to the Middle East is limited, in particular little is known about the influence of climatic factors. Mortality data from 417 shipments of sheep exported over an 11-year period (November 2004 to June 2015) were modelled retrospectively to determine associated climatic factors. The statistical analysis were performed for both the full data set with 417 voyages based on actual and estimated departure and arrival dates and a restricted data set with 71 voyages based on actual dates. The results of the full data set demonstrated a seasonal mortality pattern, with more deaths occurring on sea voyages leaving Australia in the southern hemisphere winter or spring than those departing in Australian summer or autumn. Heat stress and inadequate fat mobilisation for energy supply when sheep are inappetant on shipments may explain this seasonality. Based on these two models, the voyage and weather factors associated with sheep mortalities included departure year, autumn departure in the southern hemisphere, voyage duration, single or multiple loading port(s), weekly mean dry bulb temperature and wind speed at departure ports, and humidity at destination ports. Significant correlations were observed between weather variables at the departure ports in the Australian winter and a high sheep mortality rate during voyages. This, together with the anticipated increased heat stress risk as a result of climate change, suggests that there could be review of the trade from Australia in the southern hemisphere winter. The influence of weather at the departure ports should be considered in sheep mortality prediction models, especially Australia’s heat stress risk assessment model.

The origins and phylogeny of different sheep breeds has been widely studied using polymorphisms within the mitochondrial hypervariable region. However, little is known about the mitochondrial DNA (mtDNA) content and phylogeny based on mtDNA protein-coding genes. In this study, we assessed the phylogeny and copy number of the mtDNA in eight indigenous (population size, n=184) and three introduced (n=66) sheep breeds in China based on five mitochondrial coding genes (COX1, COX2, ATP8, ATP6 and COX3). The mean haplotype and nucleotide diversities were 0.944 and 0.00322, respectively. We identified a correlation between the lineages distribution and the genetic distance, whereby Valley-type Tibetan sheep had a closer genetic relationship with introduced breeds (Dorper, Poll Dorset and Suffolk) than with other indigenous breeds. Similarly, the Median-joining profile of haplotypes revealed the distribution of clusters according to genetic differences. Moreover, copy number analysis based on the five mitochondrial coding genes was affected by the genetic distance combining with genetic phylogeny; we also identified obvious non-synonymous mutations in ATP6 between the different levels of copy number expressions. These results imply that differences in mitogenomic compositions resulting from geographical separation lead to differences in mitochondrial function.

The purpose of the study described in this Research Communication was to report the full characterisation of the goat and sheep oxytocin-neurophysin I gene (OXT), their promoters and amino acid sequences. Using the genomic DNA as template, we sequenced and compared the whole OXT gene (3 exons), plus 958/960 nucleotides at the 5′ flanking region and 478/477 nucleotides at the 3′ flanking region, in 46 sheep and 24 goats belonging to different breeds/genetic types reared in Italy, Greece and Germany. The comparison of the obtained sequences showed a high degree of genetic variability at these loci. In particular, we focused on the SNP g.438T > C as possible example of trans-specific polymorphism. This SNP alters a putative binding site of the transcription factor Oct-1. The set-up of a luciferase assay confirmed that the C variant of this SNP negatively affects the promoter activity of the sheep OXT gene. The results of this study suggest that the SNP g.438T > C might be useful to promote association studies with traits/physiological processes controlled by this hormone.

In 1990, two selection lines of Merino sheep were established for low and high behavioural reactivity (calm and nervous temperament) at the University of Western Australia. Breeding records consistently showed that calm ewes weaned 10% to 19% more lambs than the nervous ewes. We hypothesise that calm ewes could have a higher ovulation rate than nervous ewes and/or calm ewes could have a lower rate of embryo mortality than nervous ewes. We tested these hypotheses by comparing the ovulation rate and the rate of embryo mortality between the calm and nervous lines before and after synchronisation and artificial insemination. Merino ewes from the temperament selection lines (calm, n=100; nervous, n=100) were synchronised (early breeding season) for artificial insemination (day 0) (intravaginal sponges containing fluogestone acetate and eCG immediately after sponge withdrawal). On day-17 and 11 ovarian cyclicity and corpora lutea, and on days 30 and 74 pregnancies and embryos/foetuses were determined by ultrasound. Progesterone, insulin and leptin concentrations were determined in blood plasma samples from days 5, 12 and 17. Ovarian cyclicity before and after oestrus synchronisation did not differ between the lines, but ovulation rate did (day-17: calm 1.63; nervous 1.26; P<0.01; day 11: calm 1.83; nervous 1.57; P<0.05). Ovulation rate on day 11 in nervous ewes was higher than on day-17. Loss of embryos by day 30 was high (calm: 71/150; nervous: 68/130); but nervous ewes had a lower proportion (15/47) of multiple pregnancies compared with calm ewes (30/46; P<0.01). Reproductive loss between days 30 and 74 represented 7.3% of the overall loss. Temperament did not affect concentrations of progesterone, but nervous ewes had higher insulin (32.0 pmol/l±1.17 SEM; P=0.013) and lower leptin (1.18 μg/l±0.04 SEM; P=0.002) concentrations than calm ewes (insulin: 27.8 pmol/l±1.17 SEM; leptin: 1.35 μg/l±0.04 SEM). The differences in reproductive outcomes between the calm and nervous ewes were mainly due to a higher ovulation rate in calm ewes. We suggest that reproduction in nervous ewes is compromised by factors leading up to ovulation and conception, or the uterine environment during early pregnancy, that reflect differences in energy utilisation.

Gastrointestinal nematodes represent important sources of economic losses in farmed ruminants, and the increasing frequency of anthelmintic resistance requires an increased ability to explore alternative strategies. Theoretical approaches at the crossroads of immunology and epidemiology are valuable tools in that context. In the case of Teladorsagia circumcincta in sheep, the immunological mechanisms important for resistance are increasingly well-characterized. However, despite the existence of a wide range of theoretical models, there is no framework integrating the characteristic features of this immune response into a tractable phenomenological model. Here, we propose to bridge that gap by developing a flexible modelling framework that allows for variability in nematode larval intake which can be used to track the variations in worm burdens. We parameterize this model using data from trickle infection of sheep and show that using simple immunological assumptions, our model can capture the dynamics of both adult worm burdens and nematode fecal egg counts. In addition, our analysis reveals interesting dose-dependent effects on the immune response. Finally, we discuss potential developments of this model and highlight how an improved cross-talk between empiricists and theoreticians would facilitate important advances in the study of infectious diseases.

As part of the global sheep Hapmap project, 24 individuals from each of seven indigenous Swiss sheep breeds (Bundner Oberländer sheep (BOS), Engadine Red sheep (ERS), Swiss Black-Brown Mountain sheep (SBS), Swiss Mirror sheep (SMS), Swiss White Alpine (SWA) sheep, Valais Blacknose sheep (VBS) and Valais Red sheep (VRS)), were genotyped using Illumina's Ovine SNP50 BeadChip. In total, 167 animals were subjected to a detailed analysis for genetic diversity using 45 193 informative single nucleotide polymorphisms. The results of the phylogenetic analyses supported the known proximity between populations such as VBS and VRS or SMS and SWA. Average genomic relatedness within a breed was found to be 12 percent (BOS), 5 percent (ERS), 9 percent (SBS), 10 percent (SMS), 9 percent (SWA), 12 percent (VBS) and 20 percent (VRS). Furthermore, genomic relationships between breeds were found for single individuals from SWA and SMS, VRS and VBS as well as VRS and BOS. In addition, seven out of 40 indicated parent–offspring pairs could not be confirmed. These results were further supported by results from the genome-wide population cluster analysis. This study provides a better understanding of fine-scale population structures within and between Swiss sheep breeds. This relevant information will help to increase the conservation activities of the local Swiss sheep breeds.

IgA and IgE activity against Teladorsagia circumcincta was investigated in a flock of Texel lambs following natural, mixed nematode infection among lambs. The distribution of IgA activity was similar to a gamma distribution whereas IgE activity was different. Box-Cox analysis demonstrated that X0·25 was a suitable transformation to normalise IgE responses. The transformed IgE activity was under moderate to strong genetic control. Nine different allergens were identified by proteomic analysis. Tropomyosin was selected for further analysis. IgE activity against tropomyosin was moderately heritable and associated with decreased egg counts and with reduced body weight at the time of sampling.

Understanding the frequency distribution of parasites and parasite stages among hosts is essential for efficient experimental design and statistical analysis, and is also required for the development of sustainable methods of controlling infection. Nematodirus battus is one of the most important organisms that infect sheep but the distribution of parasites among hosts is unknown. An initial analysis indicated a high frequency of animals without N. battus and with zero egg counts, suggesting the possibility of a zero-inflated distribution. We developed a Bayesian analysis using Markov chain Monte Carlo methods to estimate the parameters of the zero-inflated negative binomial distribution. The analysis of 3000 simulated data sets indicated that this method out-performed the maximum likelihood procedure. Application of this technique to faecal egg counts from lambs in a commercial upland flock indicated that N. battus counts were indeed zero-inflated. Estimating the extent of zero-inflation is important for effective statistical analysis and for the accurate identification of genetically resistant animals.

The five exons and the 5′ and 3′-untranslated regions (5′-UTR and 3′-UTR) of the oGH gene were screened for mutations using PCR-single strand conformation polymorphism (PCR-SSCP) procedures in 523 Serra da Estrela ewes and were found to be highly polymorphic. The region extending across and between the GH2-N and GH2-Z copies was sequenced allowing the design of primers for the specific PCR amplification of each copy. These were cloned and sequenced in 20 animals representative of all SSCP patterns. The corresponding genotypes were established for each copy following nucleotide sequencing of SSCP alleles. Twenty-four polymorphic sites were found at the GH2-N (or GH1) and fourteen at the GH2-Z copies. Eight amino acid substitutions were predicted at the GH2-N and six at the GH2-Z copies. Milk yield adjusted to 150 lactation days was analysed for the genotype of each oGH gene copy taken separately or together (associated genotypes) by restricted maximum likelihood (REML) through a univariate best linear unbiased prediction (BLUP) animal model with repeated measures. Significant associations between genotypes and milk yield were observed. Within GH2-N genotypes there was a milk yield differential of 21·4±0·2 l/150 d between the most (N7) and the least (N5) productive ones. Within GH2-Z genotypes there was a differential of 21·6±0·2 l/150 d between the most (Z8) and the least (Z1) productive ones. The effect of associated GH2-N and GH2-Z genotypes revealed a differential of 39·6±0·3 l/150 d between the most (N1+Z7) and the least (N3+Z2) productive associated genotypes. The results show that GH2-N and GH2-Z genotypes significantly affect milk yield in Serra da Estrela ewes. Moreover, the apparent joint effect of GH2-N and GH2-Z genotype could improve milk yield in 25% as compared with the mean milk production of the analysed population.

In ungulates, females typically need to reach a critical threshold body weight in order to reproduce. Females born in ‘poor’ years may lose 1 year of reproduction, as they reach the critical body mass 1 year later. Some studies report also a lasting effect of low initial body weight into prime age, while several other studies have shown that nearly all prime-age females ovulate or are pregnant. However, the quality of the offspring (as measured by offspring body weight) is often not considered. We tested whether female ungulates with an initially low body weight (at around weaning) reach the same reproductive output as females with higher initial weights, and thus whether or not they are able to compensate for their initial low weight. Data on body weight of 123 533 lambs of domestic sheep Ovis aries from 73 299 litters that in turn derived from 32 359 different ewes were used. All ewes and lambs had been free-ranging on outlying pastures in Norway during the entire summer season. Ewes with an initially low body weight produced smaller offspring throughout their lifespan. Ewes with an initially low body weight also produced fewer lambs at first and second parturition, but lifetime number of offspring was not related to ewes' initial weight. We concluded that bigger was better: the smaller offspring of ewes with a low initial weight demonstrate a lasting cost on reproductive effort, with consequences for animal production systems and management of wild ungulates.

Free-living Soay sheep (Ovis aries) on the island of Hirta, St Kilda, Scotland, are naturally parasitized by gastrointestinal nematodes, predominantly Teladorsagia circumcincta. In this paper we show that reduced faecal egg counts (FEC) are associated with an allele at a microsatellite locus located in the first intron of the interferon gamma gene (o(IFN)-γ) in Soay sheep lambs and yearlings, measured at approximately 4 and 16 months of age, respectively. The same allele was also associated with increased T. circumcincta-specific antibody (IgA) in lambs, but not associated significantly in yearlings. Flanking control markers failed to show a significant association with either FEC or IgA. These results suggest that a polymorphic gene conferring increased resistance to gastrointestinal nematode parasites is located at or near the interferon gamma gene, and support previous reports which have mapped a quantitative trait locus (QTL) for resistance to this region in domestic sheep. Our data are consistent with the idea that a functional polymorphism leading to reduced expression or efficacy of (IFN)-γ could enhance the immune response to gastrointestinal nematodes by favouring the activity of the Th2 cell subset and antibody associated immune mechanisms.

Domestic lamb mortality on an open forested and alpine summer range in the municipality of Målselv in northern Norway was investigated. Two hundred and fifty-three lambs in four of the area's six flocks were randomly fitted with mortality transmitters and monitored throughout the summer grazing season from June to September. Total losses in the six flocks were 183 (22.9%) lambs and 20 (4.2%) ewes. The carcasses of 69 lambs, 36 of which had radio-collars, and six ewes were recovered. Of the collared lambs, eight (22%) died as a result of disease and 27 (75%) were killed by predators. Implicated predators included red fox Vulpes vulpes, lynx Lynx lynx, golden eagle Aquila chrysaetos and wolverine Gulo gulo. Diseased animals included those diagnosed with pasteurellosis, disrupted intestinal functions and advanced coccidiosis. The remains of one lamb were too decomposed for necropsy, though there was no evidence that it had been attacked by a predator. Factors associated with losses were identified using logistic regression. Age of lamb at time of release on the summer range and age of dam were statistically associated with lamb losses (P<0.05). Mortality was higher among older lambs and those of year-old ewes. Parameters commonly associated with lamb size and growth were not statistically associated with mortality. The negative association between lamb mortality and age of dam may be related to the quality of maternal care provided by ewes of different ages, particularly vigilance in the habitat of predators. Similarly, the positive association between mortality and age of lamb may be related to changing lamb behaviour, specifically increased distance from ewe, as lambs grow older and more independent. Predators, when present, are a major cause of mortality in free-ranging lambs, as in most wild ungulates.

Twenty lambs (median body weight 37.2 kg) were kept on a paddock of 0.56 ha in England for four weeks in autumn. A 10 m grid was marked out in the paddock and from 10 of the 54 grid squares, all faeces were removed by hand at the start of each week. Faecal abundance was estimated by a point quadrat technique and observations of behaviour were made by scan sampling at intervals of 10 min during 15 h in each week. Each lamb produced on average 0.72 kg wet weight of faeces per day. Removal of faeces had no effect on grazing behaviour and the only correlation between faecal abundance and behaviour was that night lying tended to be on areas where faecal abundance was high. This was probably due to the lambs preferring certain areas for night lying and defecating heavily in these areas before starting to graze in the morning. Faecal encounter during daytime grazing was affected neither by sheep body weight nor by parasite burden. Behavioural mechanisms of avoidance of faeces are probably not detectable when behaviour is studied in the context of grid squares of 100 m2 and further study should include detailed observations on the grazing paths of individual sheep.