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As animals age they often exhibit substantial physiological and behavioural changes that can drive changes in infection status over the lifespan. Generally, immunosenescence is expected to lead to greater infection in older individuals, but this process could be confounded or counteracted by changes in behaviour, selective disappearance of certain individuals, or a range of immune processes. Here, we uncover contrasting age-related patterns of infection across three different helminth parasites in wild adult red deer (Cervus elaphus). Counts of strongyle nematodes (order: Strongylida) increased with age, while counts of liver fluke (Fasciola hepatica) and tissue worm (Elaphostrongylus cervi) decreased. These relationships could not be explained by selective disappearance of certain individuals or changes in behaviour, suggesting that immune changes could be responsible. Additionally, we found a positive relationship between social connectedness and strongyle infection, implying that previously documented age-related decreases in social behaviour may minimise exposure, reducing the negative effects of immunosenescence. These findings demonstrate that burdens of different parasites can show contrasting changes over an individual’s lifespan depending on a complex suite of intrinsic and extrinsic factors.

Helminths are common parasites of wild ungulates that can have substantial costs for growth, mortality and reproduction. Whilst these costs are relatively well documented for mature animals, knowledge of helminths' impacts on juveniles is more limited. Identifying these effects is important because young individuals are often heavily infected, and juvenile mortality is a key process regulating wild populations. Here, we investigated associations between helminth infection and overwinter survival in juvenile wild red deer (Cervus elaphus) on the Isle of Rum, Scotland. We collected fecal samples non-invasively from known individuals and used them to count propagules of 3 helminth taxa (strongyle nematodes, Fasciola hepatica and Elaphostrongylus cervi). Using generalized linear models, we investigated associations between parasite counts and overwinter survival for calves and yearlings. Strongyles were associated with reduced survival in both age classes, and F. hepatica was associated with reduced survival in yearlings, whilst E. cervi infection showed no association with survival in either age class. This study provides observational evidence for fitness costs of helminth infection in juveniles of a wild mammal, and suggests that these parasites could play a role in regulating population dynamics.

SummaryMonitoring 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 has increased across the study, and population density is known to increase parasite transmission, but we found that density 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 density than to temporal (yearly) dynamics, while abundance of coccidia showed a strong temporal trend without any influence of density. Strikingly, coccidian abundance increased threefold 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.Key findingsWe studied temporal trends in gastrointestinal parasites of wild sheep over 31 yearsYear and host population density explained temporal variation in parasites independentlyPrevalence of both strongyle nematodes and coccidia varied across the study periodAbundance of strongyles was more closely linked to host density than yearAbundance of coccidia increased threefold over time in lambs, but did not vary in adults

A longstanding prediction of life history theory states that parasitism can provide a mechanistic link between current and future fitness, thereby mediating life history tradeoffs. Demonstrating this mechanism in the wild involves linking a reproductive cost for parasitism with a cost of parasitism for fitness, which requires high-resolution longitudinal monitoring of individuals. Here, we examined associations among reproductive investment, immunity, parasitism, and fitness-related traits in a wild population of individually identified red deer (Cervus elaphus). Using path analysis, we constructed and analysed possible links among these traits, particularly investigating whether costs of lactation for survival and fecundity were likely to act through increased parasitism. Lactation was associated with increased strongyle parasitism, which was then associated with decreased subsequent fitness for all four fitness-related traits we examined. In addition, lactation was directly associated with later parturition date the following year, and increased helminth-specific antibody levels were associated with increased probability of reproduction. This study offers observational evidence for parasite mediation of multiple life history tradeoffs in a wild mammal, and supports the value of considering life history investment, immunity, and parasitism simultaneously when examining fitness consequences in disease ecology.

Helminths are common parasites of wild ungulates that can have substantial costs for growth, mortality, and reproduction. While these costs are relatively well documented for mature animals, knowledge of helminths’ impacts on juveniles is more limited. Identifying these effects is important because young individuals are often heavily infected, and juvenile mortality is an important process regulating wild populations. Here, we investigated associations between helminth infection and overwinter survival in juvenile wild red deer (Cervus elaphus) on the Isle of Rum, Scotland. We collected faecal samples non-invasively from known individuals and used them to count propagules of three helminth taxa (strongyle nematodes, Fasciola hepatica, and Elaphostrongylus cervi). Using generalised linear models, we investigated associations between parasite counts and overwinter survival for calves and yearlings. Strongyles were associated with reduced survival in both age class, and F. hepatica was associated with reduced survival in yearlings, while E. cervi infection showed no association with survival in either age class. This study provides observational evidence for fitness costs of helminth infection in juveniles of a wild mammal, and suggests that these parasites could play a role in regulating population dynamics.

International audience; Rural economies across Europe, and associated countries, heavily rely on small ruminant (SR) systems, particularly in areas unsuitable for other agriculture livestock production systems. Despite recent advances in precision livestock farming (PLF) and digital technologies (DT) to facilitate farm management, there is a slow rate of uptake by SR producers. This paper presents the project Sm@RT, which aims to improve the understanding, awareness and uptake of different technologies currently available to the SR sector, and further to facilitate ‘solutions to needs’ and identification of technology gaps. Sm@RT focus is on the use of PLF and DT across all types of sheep and goat management systems and environments, to develop a long-term, self-sustainable European/International network to encouragethe use of PLF and DT across the sectors. Eight countries (UK, Ireland, Norway, Estonia, France, Italy, Hungary, Israel) are involved in Sm@rt. The paper presents the original project’s approach: an interactive transdisciplinary and multi-actor approach relying on well-equipped demonstration farms (‘digifarms’) and innovative commercial farms to foster exchanges within the SR industry. It is an iterative step-by-step approach to engage relevant actors from all stakeholder groups at the appropriate level in the knowledge exchange process, crucial to boost dissemination in a trusting environment. Sm@RT will create opportunities to collect and exchange technical and practical knowledge, with information and guidelines on how to use PLF and DT to gain the best benefits for SR systems of production. This will increase efficiency and sustainability of SR farming systems in Europe and beyond.

1. Life history theory predicts that reproductive investment draws resources away from immunity, resulting in increased parasitism. However, studies of reproductive tradeoffs rarely examine multiple measures of reproduction, immunity, and parasitism. It is therefore unclear whether the immune costs of reproductive traits correlate with their resource costs, and whether increased parasitism emerges from weaker immunity.2. We examined these relationships in wild female red deer (Cervus elaphus) with variable reproductive investment and longitudinal data on mucosal antibody levels and helminth parasitism. We noninvasively collected faecal samples, counting propagules of strongyle nematodes (order: Strongylida), the common liver flukeFasciola hepaticaand the red deer tissue nematodeElaphostrongylus cervi. We also quantified both total and anti-strongyle mucosal IgA to measure general and specific immune investment.3. Contrary to our predictions, we found that gestation was associated with decreased total IgA but with no increase in parasitism. Meanwhile, the considerable resource demand of lactation had no further immune cost but was associated with higher counts of strongyle nematodes andElaphostrongylus cervi. These contrasting costs arose despite a negative correlation between antibodies and strongyle count, which implied that IgA was indicative of protective immunity.4. Our findings suggest that processes other than classical resource allocation tradeoffs are involved in mediating observed relationships between reproduction, immunity, and parasitism in wild mammals. In particular, reproduction-immunity tradeoffs may result from hormonal regulation or maternal antibody transfer, with parasitism increasing as a result of increased exposure arising from resource acquisition constraints. We advocate careful consideration of resource-independent mechanistic links and measurement of both immunity and parasitism when investigating reproductive costs.

The adaptive immune system is critical to an effective, long-lasting ability to respond to infection in vertebrates and T-helper (Th) cells play a key role in orchestrating the adaptive immune response. Laboratory studies show that functionally distinct Th responses provide protection against different kinds of parasites (i.e., Th1 responses against microparasites and Th2 against macroparasites).Natural populations must deal with challenges from a wide range of infectious agents and co-infection with different types of parasite is the norm, so different Th responses are likely to play an important and dynamic role in maintaining host health and fitness. However, the relationship between T helper immune phenotypes and infection with different types of parasites remains poorly understood in wild animals.In this study, we characterised variation in functionally distinct Th responses (Th1, Th2, Th17 and regulatory responses) in a wild population of Soay sheep using flow cytometry to detect Th-subset specific transcription factors, and ex vivo lymphocyte stimulation to quantify release of Th-associated cytokines. We specifically tested the prediction that raised Th1 and Th2 responses should predict reduced apicomplexan (coccidian) and helminth (nematode) parasite burdens, respectively.Cell counts of different Th subsets measured by flow cytometry did not vary with age or sex. However, all measures of Th-associated ex vivo cytokine production increased with age, and Th17- and regulatory Th-associated cytokine production increased more rapidly with age in males than females.Independent of age and sex, Th2-associated immune measures negatively predicted gastro-intestinal strongyle nematode faecal egg count, while production of the Th1-associated cytokine IFN-γ negatively predicted coccidian faecal oocyst count.Our results provide important support from outside the laboratory that Th1 and Th2 responses confer resistance to different kinds of parasites (micro- and macro-parasites, respectively). They also add to mounting evidence from wild populations that Th1/Th2 trade-offs often observed in controlled laboratory experiments may not readily translate to more complex natural systems.Our study illustrates that harnessing more specific reagents and tools from laboratory immunology has the potential to illuminate our understanding of epidemiology and host-parasite co-evolution in the wild.

Seasonality is a ubiquitous feature of wildlife disease ecology, but is determined by a complex interplay of environmental, parasitological and host factors. Gastrointestinal parasites often exhibit strong seasonal dynamics in wild vertebrate populations due to, for example, environmental influences on free-living or vectored life stages, and variation in the physiological and immune status of hosts across their annual cycle. At the same time, wild populations are typically infected with multiple parasites. The seasonal dynamics of co-infecting parasites may differ depending on age and reproductive status, and associations among parasites may be driven by short-term within-individual changes or longer-term interactions that are consistent among hosts. Here, we used faecal samples and egg counts collected repeatedly from individually marked and monitored wild Soay sheep that were part of a long-term study to investigate seasonal dynamics of six gastrointestinal parasite groups (strongyle nematodes, coccidian protozoa, Capillaria, Strongyloides, Nematodirus, and Moniezia). Prevalence and abundance generally tended to be higher spring and summer, and burdens were higher in lambs than adults. Within the highly prevalent strongyle nematode group, we found differences in seasonality of egg counts depending on adult reproductive status. Reproductive ewes had increased counts in spring around the time of birth followed by a drop in abundance in summer, while barren ewes showed little evidence of seasonality. Males showed a sustained rise in egg counts through spring and summer, and sex differences were only strongly apparent in summer. In contrast, in similarly prevalent coccidia we found a peak in faecal oocyst counts in spring but no differences in seasonality among males, barren and pregnant ewes. Using multivariate mixed-effects models, we went on to show that both strongyle and coccidia counts are moderately repeatable across seasons among individuals. We further show that apparent positive correlation between strongyle and coccidia counts was driven by short-term within-individual changes in both parasite burdens rather than long-term among-individual covariation. Overall, our results demonstrate that seasonality varies across demographic and parasite groups and highlight the value of investigating fluctuating susceptibility and exposure over time for understanding epidemiology of a population.

Refugia-based strategies are intended to help slow the development of anthelmintic resistance by providing a population of parasites that are not exposed to the treatment. Evidence from field studies is lacking. There is no single way to incorporate refugia into a parasite control program. There are many options available varying greatly in complexity and practicality, and none are suitable for all situations. Incorporating refugia into production systems typically requires a change in farmer mindset and greater input of time, labor, and/or technology, but is necessary to help preserve anthelmintic efficacy and promote sustainable animal production systems.

The Teladorsagia circumcincta P-glycoprotein-9 (Tci-pgp-9) gene has previously been implicated in multiple-anthelmintic resistance in this parasite. Here we further characterise genetic diversity in Tci-pgp-9 and its possible role in ivermectin (IVM) and multi-drug resistance using two UK field isolates of T. circumcincta, one susceptible to anthelmintics (MTci2) and the other resistant to most available anthelmintics including IVM (MTci5). A comparison of full-length Tci-pgp-9 cDNA transcripts from the MTci2 and MTci5 isolates (∼3.8 kb in both cases) indicated that they shared 95.6% and 99.5% identity at the nucleotide and amino acid levels, respectively. Nine non-synonymous SNPs were found in the MTci5 sequences relative to their MTci2 counterparts. Twelve genomic sequence variants of the first internucleotide binding domain of Tci-pgp-9 were identified and up to 10 of these were present in some individual worms, strongly supporting previous evidence that amplification of this gene has occurred in T. circumcincta. On average, fewer distinct sequence variants of Tci-pgp-9 were present in individual worms of the MTci5 isolate than in those of the MTci2 isolate. A further reduction in the number of sequence variants was observed in individuals derived from an IVM-treated sub-population of MTci5. These findings suggest that Tci-pgp-9 was under purifying selection in the face of IVM treatment in T. circumcincta, with some sequence variants being selected against., Graphical abstract Image 1, Highlights • Tci-pgp-9 cDNA sequences from the MTci2 and MTci5 isolates shared 95.6% identity. • Comparison of Tci-pgp-9 sequences from MTci5 and MTci2 showed 9 non-synonymous SNPs. • Multiple Tci-pgp-9-IBDA sequence variants occurred in the majority of individual larvae. • Tci-pgp-9 appears to be under purifying selection after anthelmintics are applied.

Although mountain sheep systems suffer from climatic and environmental handicaps that constrain productivity and economic viability, they have an important economic role, maintain habitats and species of high nature conservation value and support the provision of a range of ecosystem services of benefit to society. Using Precision Livestock Farming (PLF) in extensive mountain sheep systems could bring benefits for animal performance, economical performance and labour. This paper presents results from a 3 year experiment where PLF principles were implemented on an extensive mountain sheep farm and an assessment made of whether or not such an approach could benefit more marginal sheep systems. A 900 ewe flock (600 Scottish Blackface ewes, 300 Lleyn ewes) was divided equally into two separate systems, one where the flock was managed conventionally (CON) at group level, and the other where the individuals in the flock were subjected to a PLF management protocol where electronic weighing, recording and drafting equipment were used, linked to the electronic identification (EID) tags of the animals. Two main management strategies were compared and contrasted; one relating to winter feeding of the pregnant ewes, the other relating to anthelmintic treatment of lambs during the summer. Yearly labour profiles were created by measuring the time spent doing individual tasks associated with the two management systems. Net margins (£/ewe) were calculated for the two systems. Additionally, the yearly labour profiles were scaled-up using commercial data to quantify potential labour savings on more traditionally managed mountain farms if PLF principles were adopted. Analyses indicated that the two different management systems did not result in any significant difference in terms of ewe weights, mid-pregnancy scanning figures, ewe and lamb mortality rates, or lamb weight post-weaning. However, the proportion of lambs needing anthelmintic treatment was significantly reduced by 40% between the CON and the PLF, resulting in a reduction of 46% in the amount of anthelmintic used. Over a whole year, the total amount of labour required in the PLF management system was reduced by 36%. Across the 3 years, the net margin for the two systems showed an average difference of £3/ewe higher in the PLF. For a more traditional farm embracing a PLF approach, analyses suggested labour reduction of 19%, equating to £1.60/ewe savings. This study shows that it is beneficial for farmers to consider managing a mountain ewe flock at an individual rather than at flock or batch level using PLF technology.

International audience; Tae benefit of precision livestock fanning (PLF) is well recognised in the more intensive livestock sec.tors, such as dairy, pig and poultry. However, PLF has not been applied as \videly in species where animals are con5idered to have a lower individual value or with Jess ec.onomic interest, as is the case in small ruminants (SR), or in extensive management systems. This is despite the very significant production, welfare, and labour efficiency advantages that can be achieved by applying PLF in these contexts. Despite their crucial role for the rural economy, society, and environment and their importance in ecosystem services such as biodiversity and maintaining cultural heritage, SR systems face issues such as challenging clinlatic and topographie.al conditions, Jack of labour and low profitability that could be alleviated by introducing PLF technologies. Research on PLF for systems bas been recently emerging, but perhaps Jacks a joint up approach. Tiris paper present an overview of current research and potential for future applications in several countries of PLF for SR system5 on varions themes, highlighting the wealth of potentially available solutions and prototypes. The topics presented cover feed and water intake, health, disease and parasite control and mo1ritoring, fertility and reproduction management, grazing and predation control, animal locations and management monitoring, lambing and mismothering issues, as well as lactation monitoring. Issues relating to acceptability, economic relevance, technology readiness level (TRL) and industry engagement will also be discussed.

Anthelmintic resistance is a threat to global food security. In order to alleviate the selection pressure for resistance and maintain drug efficacy, management strategies increasingly aim to preserve a proportion of the parasite population in ‘refugia’, unexposed to treatment. While persuasive in its logic, and widely advocated as best practice, evidence for the ability of refugia-based approaches to slow the development of drug resistance in parasitic helminths is currently limited. Moreover, the conditions needed for refugia to work, or how transferable those are between parasite-host systems, are not known. This review, born of an international workshop, seeks to deconstruct the concept of refugia and examine its assumptions and applicability in different situations. We conclude that factors potentially important to refugia, such as the fitness cost of drug resistance, the degree of mixing between parasite sub-populations selected through treatment or not, and the impact of parasite life-history, genetics and environment on the population dynamics of resistance, vary widely between systems. The success of attempts to generate refugia to limit anthelmintic drug resistance are therefore likely to be highly dependent on the system in hand. Additional research is needed on the concept of refugia and the underlying principles for its application across systems, as well as empirical studies within systems that prove and optimise its usefulness., Graphical abstract Image 1, Highlights • Refugia is currently recommended for strategic control of AR in GI nematodes of livestock. • Conditions required for refugia-based control may differ between parasite systems. • An improved understanding of the parameters influencing refugia-based control is required.

Spatial heterogeneity in parasite susceptibility and exposure is a common source of confounding variation in disease ecology studies. However, it is not known whether spatial autocorrelation acts on immunity in particular at small scales, within wild animal populations, and whether this predicts spatial patterns in infection. Here we used a well-mixed wild population of individually recognised red deer (Cervus elaphus) inhabiting a heterogeneous landscape to investigate fine-scale spatial patterns of immunity and parasitism. We noninvasively collected 842 faecal samples from 141 females with known ranging behaviour over two years. We quantified total and helminth-specific mucosal antibodies and counted propagules of three gastrointestinal helminth taxa. These data were analysed with linear mixed models using the Integrated Nested Laplace Approximation (INLA), using a Stochastic Partial Differentiation Equation approach (SPDE) to control for and quantify spatial autocorrelation. We also investigated whether spatial patterns of immunity and parasitism changed seasonally. We discovered substantial spatial heterogeneity in general and helminth-specific antibody levels and parasitism with two helminth taxa, all of which exhibited contrasting seasonal variation in their spatial patterns. Notably, strongyle nematode intensity did not align with density hotspots, while Fasciola hepatica intensity appeared to be strongly influenced by the presence of wet grazing. In addition, antibody hotspots did not correlate with distributions of any parasites. Our results suggest spatial heterogeneity may be an important factor affecting immunity and parasitism in a wide range of study systems. We discuss these findings with regards to the design of sampling regimes and public health interventions, and suggest that disease ecology studies investigate spatial heterogeneity more regularly to enhance their results, even when examining small geographic areas.

Molecular methods based on ITS2 sequence analysis were used to identify strongylid parasites and describe their diversity in a management intervention and anthelmintic drug treatment-free sheep flock. Fourteen different nematode parasite species were identified in the flock and the results showed a greater level of nematode species diversity than is normally reported in managed farmed flocks, with the presence of parasites such as Bunostomum trigonocephalum, Ostertagia leptospicularis, Spiculopteragia houdemeri and Trichostrongylus retortaeformis that are considered to be absent or rare in sheep kept in comparable localities. The implied prevalences of Haemonchus contortus in lambs, and of Trichostrongylus axei in lambs, ewes and rams, were higher than those in farmed sheep kept in similar regions, while those of Teladorsagia circumcincta and Trichostrongylus vitrinus were lower. Comparison of the patterns of nematode parasite infection between the summer and autumn sampling periods showed differences from the scenarios that are commonplace in comparable managed flocks; with T. vitrinus burdens of the lambs being higher in the summer than in the winter, and Oesophagostomum venulosum being the predominant nematode species in the adult sheep during the summer, while more-or-less absent from these groups during the winter. Rams played an important role in the epidemiology of certain parasitic nematode species. The relatively non-pathogenic O. venulosum was the only parasitic nematode species to predominate in any group during the study. This preliminary characterisation of the nematode parasite burdens of sheep extensively grazed on diverse unimproved pastures will aid in the understanding of the parasitological consequences of intensive grazing management and of the manner in which modern agriculture upsets the equilibrium between parasites and their hosts. These factors must be accounted for when defining the concept of sustainable parasite control and informing sustainability with reference to commercially efficient sheep farming.

In small ruminants, faecal egg counts (FECs) and reduction in FECs (FECR) are the most common methods for the assessment of intensity of gastrointestinal (GI) nematodes infections and anthelmintic drug efficacy, respectively. The main limitation of these methods is the time and cost to conduct FECs on a representative number of individual animals. A cost-saving alternative would be to examine pooled faecal samples, however little is known regarding whether pooling can give representative results. In the present study, we compared the FECR results obtained by both an individual and a pooled examination strategy across different pool sizes and analytical sensitivity of the FEC techniques. A survey was conducted on 5 sheep farms in Scotland, where anthelmintic resistance is known to be widespread. Lambs were treated with fenbendazole (4 groups), levamisole (3 groups), ivermectin (3 groups) or moxidectin (1 group). For each group, individual faecal samples were collected from 20 animals, at baseline (D0) and 14 days after (D14) anthelmintic administration. Faecal samples were analyzed as pools of 3-5, 6-10, and 14-20 individual samples. Both individual and pooled samples were screened for GI strongyle and Nematodirus eggs using two FEC techniques with three different levels of analytical sensitivity, including Mini-FLOTAC (analytical sensitivity of 10 eggs per gram of faeces (EPG)) and McMaster (analytical sensitivity of 15 or 50 EPG).For both Mini-FLOTAC and McMaster (analytical sensitivity of 15 EPG), there was a perfect agreement in classifying the efficacy of the anthelmintic as 'normal', 'doubtful' or 'reduced' regardless of pool size. When using the McMaster method (analytical sensitivity of 50 EPG) anthelmintic efficacy was often falsely classified as 'normal' or assessment was not possible due to zero FECs at D0, and this became more pronounced when the pool size increased. In conclusion, pooling ovine faecal samples holds promise as a cost-saving and efficient strategy for assessing GI nematode FECR. However, for the assessment FECR one will need to consider the baseline FEC, pool size and analytical sensitivity of the method.

Parasitism in wild mammals can vary according to myriad intrinsic and extrinsic factors, many of which vary seasonally. However, seasonal variation in parasitism is rarely studied using repeated samples from known individuals. Here we used a wild population of individually recognized red deer (Cervus elaphus) on the Isle of Rum to quantify seasonality and intrinsic factors affecting gastrointestinal helminth parasitism over the course of a year. We collected 1020 non-invasive faecal samples from 328 known individuals which we then analysed for propagules of three helminth taxa: strongyle nematodes, the common liver fluke Fasciola hepatica and the tissue nematode Elaphostrongylus cervi. Zero-inflated Poisson models were used to investigate how season, age and sex were associated with parasite prevalence and count intensity, while Poisson models were used to quantify individual repeatability within and between sampling seasons. Parasite intensity and prevalence varied according to all investigated factors, with opposing seasonality, age profiles and sex biases between parasite taxa. Repeatability was moderate, decreased between seasons and varied between parasites; both F. hepatica and E. cervi showed significant between-season repeatability, while strongyle nematode counts were only repeatable within-season and showed no repeatability within individuals across the year.

Parasitic roundworm infections are ubiquitous in grazing livestock. Chemical control through the frequent 'blanket' administration of anthelmintics (wormers) has been, and remains, the cornerstone in controlling these infections, but this practice is unsustainable. Alternative strategies are available but, even with the plethora of best practice advice available, have yet to be integrated into routine farming practice. This is probably due to a range of factors, including contradictory advice from different sources, changes to advice following increased scientific understanding, and top-down knowledge exchange patterns. In this article, we discuss the worm control options available, the translation of new best practice advice from science bench to field, and ideas for future work and directions.

Global agriculture will be required to intensify production from a shrinking natural resource base. Helminth infections of ruminants are a major constraint on efficient livestock production. The current challenge is to develop diagnostic methods that detect the production impact of helminth infections on farms in order to target control measures and contribute to the global challenge of preserving food security. We review here our understanding of the effects of helminth infections and control practices on productivity and the diagnostic tools that can inform on this. By combining advances in helminth laboratory diagnostics and animal health economics, sustainable management of helminth infections can be integrated into the whole-farm economic context.

The sustainability of sheep production is hindered by anthelmintic resistance. Options to slow down or prevent resistance have been widely studied but their application in the field is still limited. In this study, the practical application and effect of a targeted selective treatment (TST) approach for the treatment of parasitic gastroenteritis was investigated in lambs (n = 385) over a 2 year period. At 14-day intervals during the grazing season, liveweight, breech soiling and anthelmintic treatments were individually recorded. Selection of lambs for anthelmintic treatment in the TST group was based on pre-calculated individual growth rates, with a matched cohort routinely treated (RT) with anthelmintic drug every 6 weeks. The adoption of a TST approach had no negative effect on the liveweight gains of the lambs, time to finishing or breech soiling measures compared to RT lambs; however a 50% decrease in anthelmintic treatment was observed in the TST group. The time to implement this system averaged 2 min per lamb. It is concluded that the TST could be suitable for commercial sheep farms, in association with automated weighing systems, potentially reducing selection for anthelmintic resistance, while having no negative effect on production.

A major constraint on the effective control and management of helminth parasites in livestock is the lack of rapid and reliable diagnostic tests to identify the parasite species responsible for disease and to allow informed treatment decisions to be made. In the present study, we have developed a novel DNA-based assay for the detection of Haemonchus contortus eggs in ovine faecal samples, using loop-mediated isothermal amplification (or LAMP). LAMP allows for rapid detection of H. contortus DNA under isothermal incubation conditions. The robust nature of this assay negates the need for extensive DNA extraction, allowing amplification from relatively crude samples. Preliminary results suggest that LAMP is highly specific, and does not cross-react with DNA from other common co-infecting parasites. The Haemonchus LAMP assay is also highly sensitive, exhibiting a 10 times lower detection limit than the equivalent PCR; 10−5 ng/μl and 10−4 ng/μl DNA, respectively, allowing detection in a faecal samples containing two Haemonchus eggs per gram. Translation of this assay onto a real-time platform provided rapid results and highlighted its potential as a quantitative assay which could inform treatment decisions in the future.

Sustainable control of nematode parasites in small ruminant production is a worldwide ambition. Development of anthelmintic resistance can severely impair small ruminant production. A practical approach to reduce selection pressure for anthelmintic resistance is to treat only a proportion of the flock (Targeted Selective Treatment), leaving a proportion of the nematode population untreated. The aim of this study was to compare the sustainability and efficacy of a performance-based marker, the Happy Factor™, a monitor of nutrient utilisation efficiency, with a routine whole flock anthelmintic treatment. In a commercial flock in the South West of Scotland, 183 Texel cross lambs were split into two matched but co-grazing groups: one group managed as routinely for the farm (RT group) and the other subjected to targeted selective treatment (TST group). All lambs from the RT group were drenched every 6 weeks during the grazing season, while anthelmintic administration in the TST group was restricted to animals that failed to reach pre-determined weight gain targets, based on an estimate of their efficiency of gross energy utilisation. Animal performance and parasitological data were recorded every two weeks. In the 20 week period of the study, anthelmintic treatments were reduced by approximately 50% in the TST group compared to a routine anthelmintic administration that would have been applied, whilst epg counts were always

Infections with parasitic helminths (nematodes and trematodes) represent a significant economic and welfare burden to the global ruminant livestock industry. The increasing prevalence of anthelmintic resistance means that current control programmes are costly and unsustainable in the long term. Recent changes in the epidemiology, seasonality and geographic distribution of helminth infections have been attributed to climate change. However, other changes in environment (e.g., land use) and in livestock farming, such as intensification and altered management practices, will also have an impact on helminth infections. Sustainable control of helminth infections in a changing world requires detailed knowledge of these interactions. In particular, there is a need to devise new, sustainable strategies for the effective control of ruminant helminthoses in the face of global change. In this paper, we consider the impact of helminth infections in grazing ruminants, taking a European perspective, and identify scientific and applied priorities to mitigate these impacts. These include the development and deployment of efficient, high- throughput diagnostic tests to support targeted intervention, modelling of geographic and seasonal trends in infection, more thorough economic data and analysis of the impact of helminth infections and greater translation and involvement of end-users in devising and disseminating best practices. Complex changes in helminth epidemiology will require innovative solutions. By developing and using new technologies and models, the use of anthelmintics can be optimised to limit the development and spread of drug resistance and to reduce the overall economic impact of helminth infections. This will be essential to the continued productivity and profitability of livestock farming in Europe and its contribution to regional and global food security. View Full-Text

Background: Refugia based anthelmintic protocols aim to reduce the rate of development of anthelmintic resistance in gastrointestinal nematodes (GIN). Previous studies have illustrated the impact of different drenching regimes on drug efficacy and animal growth; however, the impact on nematode populations has yet to be characterised within natural infections. This study investigated the changes in species composition of GIN throughout the grazing season, following implementation of four different ivermectin drenching regimes over six years: neo-suppressive monthly treatment (NST), targeted selective treatment (TST), strategic prophylactic treatment (SPT) and treatment upon observation of clinical signs (MT).Methods: Lambs were grazed on one of eight replicate paddocks each grazing season following treatment regimes assigned in year 1. Faecal samples were collected fortnightly from all animals and hatched to first stage larvae (L1). DNA was extracted from individual L1 and a multiplex PCR assay targeting the internal transcribed spacer 2 (ITS2) region of Teladorsagia circumcincta, Trichostrongylus spp. and Haemonchus contortus conducted. Other species were identified using species-specific PCR. Worm-naïve tracer lambs were grazed on the paddocks at the start and end of each grazing season and adult worms recovered at post mortem to investigate the parasite population on pasture.Results: Results showed an overall decrease in species diversity in egg output from the NST group which occurred within a single grazing season and was consistent throughout the experiment. Species diversity was protected over six years in groups implementing TST, SPT and MT treatment regimes, designed to offer refugia. The expected shift in species prevalence throughout the season from Teladorsagia to Trichostrongylus was observed in all but the NST group where only Teladorsagia spp. were recovered from trial lambs by the end of the experiment. Worm burdens indicated the presence of several species at relatively low abundance on pasture in the NST group in 2011. However, these species were not represented in egg output from trial lambs, probably due to the frequent anthelmintic treatment administered throughout the grazing season.Conclusion: The molecular methods utilised here worked well. The comparable results of the three refugia-based treatment regimes suggest that nematode diversity can be maintained using part or whole group treatments if a rich supra-population of parasites are available to re-infect animals post treatment.

The aim of this study was to examine the use of Happy Factor™ weight based targeted selective treatment (TST) on several commercial farms in Scotland in combination with findings from a long term trial on a research farm to assess the potential for TST use in varying farming operations as an alternative to the current regimen of whole flock treatment. Lambs on each farm were regularly weighed and climatic conditions and pasture availability measured for inclusion into the Happy Factor™ model to calculate weight targets. Half of the lambs were allocated to TST treatment and any failing to reach the weight target was treated with the anthelmintic of choice on that farm, while the remaining half of each flock was treated with anthelmintic as per normal practice on that farm (routine treatment, RT). The research farm (farm 1) hosted a long term trial using four anthelmintic treatment regimes over 6 years, and data from two regimes are presented here, alongside findings from three further farms: two commercial enterprises (farms 2 and 3) and a research farm operating as a commercial analogue with two breeds (farms 4a and 4b). The effect of TST strategy on lamb productivity and the number of anthelmintic treatments was investigated. There was no evidence (p>0.300) that mean bodyweight or growth rate was different between TST and RT groups on any of the farms and 95% confidence intervals of TST and RT groups generally suggested that TST had negligible unfavourable effects on the average growth of lambs for most of the farms. Growth rates ranged from 97.39 to 189.16g/day reflecting the varied nature of the farms. All commercial farms used significantly less (1.34 RT versus 1.14 TST treatments per animal, p

Malaria in eastern Sudan is characterised by limited seasonal transmission, with the majority of the year remaining transmission-free. Some inhabitants who contract malaria during the transmission season retain long-lasting sub-patent infections, which probably initiate transmission the following year. Here we have monitored Plasmodium falciparum infection prevalence and gametocyte production during the dry season, and examined the impact of parasite genetic multiplicity on infection longevity. A cohort of 38 individuals who were infected with P. falciparum in November 2001 was monitored monthly by microscopy and PCR until December 2002. Reverse transcriptase polymerase chain reaction of the pfg377 gene was used to detect sub-patent gametocytes. In addition, all isolates were examined for msp-2 alleles and the mean number of parasite clones per infection was estimated. We found that a large proportion (40%) of the cohort retained gametocytes throughout the dry season. The majority of patients retained asexual infection for at least 7 months. Genetic multiplicity of P. falciparum significantly influenced longevity of asexual infection and its gametocyte production. Gametocytes from mixed genotype P. falciparum infections persisted three times longer than those from single genotype infections, suggesting that genetic diversity promotes persistence. These findings are discussed in the context of the parasite biology and malaria epidemiology in the study area.

The sheep scab mite, Psoroptes ovis, causes severe dermatitis in infected sheep with severe welfare and production implications. The dermatitis has the characteristics of an immediate hypersensitivity type reaction which, by analogy to other mite species, including the house dust mites (Dermatophagoides spp.), is likely to be invoked by a variety of allergens including mite-derived proteinases. Here, the proteinases in P. ovis extracts were characterised using substrate gel analysis, inhibitor sensitivity and their ability to degrade a variety of potential natural protein substrates. These analyses showed that mites contain several proteinases which could be differentiated on the basis of molecular size and inhibitor sensitivity with cysteine, metalloproteinases and aspartyl proteinases predominating. These proteinases degraded collagen and fibronectin, possibly indicative of a role in lesion initiation, they degraded several blood proteins, a property which may aid mite feeding and they degraded immunoglobulin G, possibly aiding immuno-evasion. Because proteinases, particularly the cysteine class, are demonstrably allergenic in other mite infestations, these proteinases clearly merit further immunological and biochemical definition.

In the UK, recent mean temperatures have consistently increased by between 1°C and 4°C compared to the 30-year monthly averages. Furthermore, all available predictive models for the UK indicate that the climate is likely to change further and feature more extreme weather events and a trend towards wetter, milder winters and hotter, drier summers. These changes will alter the prevalence of endemic diseases spatially and/or temporally and impact on animal health and welfare. Most notable among these endemic parasites are the helminths, which have been shown to be very strongly influenced by both the short-term weather and climate through effects on their free-living larval stages on pasture. In this review, we examine recent trends in prevalence and epidemiology of key helminth species and consider whether these could be climate-related. We identify likely effects of temperature and rainfall on the free-living stages and some key parasite traits likely to determine parasite abundance under changed climatic conditions. We find clear evidence that climate change, especially elevated temperature, has already changed the overall abundance, seasonality and spatial spread of endemic helminths in the UK. We explore some confounders and alternative explanations for the observed patterns. Finally, we explore the implications of these findings for policy makers and the livestock industry and make some recommendations for future research priorities.

Background Sheep scab is a highly contagious disease of sheep caused by the ectoparasitic mite Psoroptes ovis. The disease is endemic in the UK and has significant economic impact through its effects on performance and welfare. Diagnosis of sheep scab is achieved through observation of clinical signs e.g. itching, pruritis and wool loss and ultimately through the detection of mites in skin scrapings. Early stages of infestation are often difficult to diagnose and sub-clinical animals can be a major factor in disease spread. The development of a diagnostic assay would enable farmers and veterinarians to detect disease at an early stage, reducing the risk of developing clinical disease and limiting spread. Methods Serum samples were obtained from an outbreak of sheep scab within an experimental flock (n = 480 (3 samples each from 160 sheep)) allowing the assessment, by ELISA of sheep scab specific antibody prior to infestation, mid-outbreak (combined with clinical assessment) and post-treatment. Results Analysis of pre-infestation samples demonstrated low levels of potential false positives (3.8%). Of the 27 animals with clinical or behavioural signs of disease 25 tested positive at the mid-outbreak sampling period, however, the remaining 2 sheep tested positive at the subsequent sampling period. Clinical assessment revealed the absence of clinical or behavioural signs of disease in 132 sheep, whilst analysis of mid-outbreak samples showed that 105 of these clinically negative animals were serologically positive, representing potential sub-clinical infestations. Conclusions This study demonstrates that this ELISA test can effectively diagnose sheep scab in a natural outbreak of disease, and more importantly, highlights its ability to detect sub-clinically infested animals. This ELISA, employing a single recombinant antigen, represents a major step forward in the diagnosis of sheep scab and may prove to be critical in any future control program.

Background Sheep scab is caused by Psoroptes ovis and is arguably the most important ectoparasitic disease affecting sheep in the UK. The disease is highly contagious and causes and considerable pruritis and irritation and is therefore a major welfare concern. Current methods of treatment are unsustainable and in order to elucidate novel methods of disease control a more comprehensive understanding of the parasite is required. To date, no full genomic DNA sequence or large scale transcript datasets are available and prior to this study only 484 P. ovis expressed sequence tags (ESTs) were accessible in public databases. Results In order to further expand upon the transcriptomic coverage of P. ovis thus facilitating novel insights into the mite biology we undertook a larger scale EST approach, incorporating newly generated and previously described P. ovis transcript data and representing the largest collection of P. ovis ESTs to date. We sequenced 1,574 ESTs and assembled these along with 484 previously generated P. ovis ESTs, which resulted in the identification of 1,545 unique P. ovis sequences. BLASTX searches identified 961 ESTs with significant hits (E-value < 1E-04) and 584 novel P. ovis ESTs. Gene Ontology (GO) analysis allowed the functional annotation of 880 ESTs and included predictions of signal peptide and transmembrane domains; allowing the identification of potential P. ovis excreted/secreted factors, and mapping of metabolic pathways. Conclusions This dataset currently represents the largest collection of P. ovis ESTs, all of which are publicly available in the GenBank EST database (dbEST) (accession numbers FR748230 - FR749648). Functional analysis of this dataset identified important homologues, including house dust mite allergens and tick salivary factors. These findings offer new insights into the underlying biology of P. ovis, facilitating further investigations into mite biology and the identification of novel methods of intervention.

All donor blood samples must be tested pretransfusion to determine the donor blood type. Standard testing protocols require that assays be performed for important bloodborne pathogens such as hepatitis C, syphilis, hepatitis B, and human immunodeficiency virus. We have demonstrated proof of the concept that a protein microarray can type whole blood and detect antibody to significant pathogens simultaneously from the same donor blood sample. The data collected demonstrate the ability of the array to accurately type blood samples while also detecting the presence of antibodies against both human immunodeficiency virus and hepatitis C virus. In conclusion, we have successfully developed a platform capable of typing human whole blood samples, while at the same time testing for the presence of antibodies specific for human immunodeficiency virus/hepatitis C virus. The major benefits of this system are its amenability to expansion with additional assays, for example, rhesus typing and syphilis and/or hepatitis B virus detection, and also the adaptability of the assay to higher-throughput analysis, currently 16 individual samples per slide, but readily expandable to a 96-well format. (C) 2008 Elsevier Inc. All rights reserved.

Serum from successful vaccine trials against the sheep scab mite, Psoroptes ovis, was used to immunoscreen a cDNA library constructed from mixed-stage and gender P. ovis to identify potential recombinant vaccine candidates. Immunodominant recombinant proteins recognised by IgG in these sera were selected for further analysis. Two candidates were identified in this way; a catchin-like protein (CLP) and a novel mu class glutathione S-transferase (GST). Both candidates were expressed in bacteria as recombinant proteins, the GST as an active enzyme, and combined with four other recombinant allergens in a multi-component recombinant vaccine. Strong serum IgG responses were induced in sheep against each of the components of the recombinant vaccine, however, the protective efficacy of the vaccine could not be determined because of variability in the establishment of a challenge infection.

Background Sheep scab is caused by the ectoparasitic mite Psoroptes ovis which initiates a profound cutaneous inflammatory response, leading to the development of the skin lesions which are characteristic of the disease. Existing control strategies rely upon injectable endectocides and acaricidal dips but concerns over residues, eco-toxicity and the development of acaricide resistance limit the sustainability of this approach. In order to identify alternative means of disease control, a deeper understanding of both the parasite and its interaction with the host are required. Methods Herein we describe the development and utilisation of an annotated P. ovis cDNA microarray containing 3,456 elements for the measurement of gene expression in this economically important ectoparasite. The array consists of 981 P. ovis EST sequences printed in triplicate along with 513 control elements. Array performance was validated through the analysis of gene expression differences between fed and starved P. ovis mites. Results Sequences represented on the array include homologues of major house dust mite allergens and tick salivary proteins, along with factors potentially involved in mite reproduction and xenobiotic metabolism. In order to validate the performance of this unique resource under biological conditions we used the array to analyse gene expression differences between fed and starved P. ovis mites. These analyses identified a number of house dust mite allergen homologues up-regulated in fed mites and P. ovis transcripts involved in stress responses, autophagy and chemosensory perception up-regulated in starved mites. Conclusion The P. ovis cDNA microarray described here has been shown to be both robust and reproducible and will enable future studies to analyse gene expression in this important ectoparasite.