Differences within Churra breed sheep in the early immune response to the infection by Teladorsagia circumcincta.

This study describes early immunological mechanisms that underlie resistance to Teladorsagia circumcincta infection in adult Churra sheep. After a first experimental infection, 6 animals were classified as resistant (RG) and 6 as susceptible (SG) to T. circumcincta infection based on their cumulative faecal egg count (cFEC) at the end of the infection. RG showed higher IgA levels against somatic antigen of T. circumcincta fourth-larvae stage (L4) in serum at day 3 post-infection (pi) (p < 0.05) and close to significance at day 21 pi (p = 0.06). Moreover, a strong negative correlation between cFEC and specific IgA was only significant in RG at day 3 pi (r = - 0.870; p < 0.05), but absent in SG. At the end of this infection, sheep were treated with moxidectin and infected again 3 weeks later to be slaughtered at day 7 pi. At necropsy, the specific IgA levels in gastric mucosa were similar between groups; the absence differences at day 7 pi could be due to a previous increase in the IgA response, probably around day 3 pi, as described during the first infection. L4 burden, 68% lower in RG than in SG, was influenced by the specific IgA in gastric mucus and the number of γδ T cells. RG group showed a positive correlation between γδ T cells and eosinophils (r = 0.900; p = 0.037); however, this correlation was not found in SG. These results show that these two phenotypes show different early immune response pattern to T. circumcincta infection in Churra sheep.

Host immunity shapes the impact of climate changes on the dynamics of parasite infections.

Global climate change is predicted to alter the distribution and dynamics of soil-transmitted helminth infections, and yet host immunity can also influence the impact of warming on host-parasite interactions and mitigate the long-term effects. We used time-series data from two helminth species of a natural herbivore and investigated the contribution of climate change and immunity on the long-term and seasonal dynamics of infection. We provide evidence that climate warming increases the availability of infective stages of both helminth species and the proportional increase in the intensity of infection for the helminth not regulated by immunity. In contrast, there is no significant long-term positive trend in the intensity for the immune-controlled helminth, as immunity reduces the net outcome of climate on parasite dynamics. Even so, hosts experienced higher infections of this helminth at an earlier age during critical months in the warmer years. Immunity can alleviate the expected long-term effect of climate on parasite infections but can also shift the seasonal peak of infection toward the younger individuals.

Carbon dioxide is an absolute requirement for exsheathment of some, but not all, abomasal nematode species.

The first step in the infection process of grazing ruminants by gastrointestinal nematodes is the exsheathment of the third-stage larva (L3). Exsheathment of various species can be achieved in vitro using carbon dioxide (CO2) under the appropriate temperature and pH conditions. However, it remains unclear whether elevated CO2 levels are an absolute requirement for exsheathment. Exsheathment of four abomasal species was investigated in both the presence and absence of CO2, in either rumen fluid (cow or sheep) or buffer (standard or enriched). Exsheathment of Ostertagia ostertagi, Teladorsagia circumcincta and Ostertagia leptospicularis was observed in CO2-depleted rumen fluid and enriched buffer (respectively 46%, 22% and 15% in rumen fluid and 28% 18% and 26% in enriched buffer after 24 h). The level of this response was dependent on the species as well as the medium, and exsheathment was significantly higher in the presence of CO2. For Haemonchus contortus, exsheathment could only be achieved under CO2-saturated conditions. In conclusion, even though these parasite species exsheath in the same environment, there were significant differences in the minimal requirements to trigger their exsheathment. Some abomasal species were capable of exsheathment in the absence of CO2, which is likely facilitated by cofactors present in the rumen fluid and/or enriched buffer.

Effects of selected Palestinian plants on the in vitro exsheathment of the third stage larvae of gastrointestinal nematodes.

BACKGROUND: Gastrointestinal parasites are one of the main restrictions to small ruminant production. Their pathological importance is primarily related to the major production losses, in quantity or quality, induced by the direct action of worms. Control of these parasites is based exclusively on the frequent use of anthelmintic drugs. However, the resistance to anthelmintics in worm populations after commercialisation of chemical drugs is now widespread. Therefore, there is a need to find new natural resources to ensure sustainable and effective treatment and control of these parasites. The aim of this study was to evaluate the anthelmintic activity, as minimum inhibitory concentration (IC 50 mg/mL), of different plant extracts using larval exsheathment inhibition assay using a two-species but steady population of parasitic nematodes (ca. 20% Teladorsagia circumcinta and 80% Trichostrongylus colubriformis). RESULTS: The study showed that the ethanolic extracts of 22 out of the 48 plant extracts, obtained from 46 plant species, have an inhibitory effect >50% (at concentrations of 100 mg/mL) on the third stage larvae (L3) of the nematodes exhibited the strongest inhibition activity (94%) with IC 50 of 0.02 mg/mL, where other members of the Rhamnaceae family have shown to possess strong anthelmintic activity (70-89%). CONCLUSIONS: Plant extracts are potential rich resources of anthelmintics to combat helminthic diseases. Our results suggest that extracts from Rhamnus elaternus, Epilobium hirsutum, Leucaena leucocephala and Rhamnus palaestinus have promising anthelmintic activity, with potential applications in animal therapeutics and feed.

Rapid method for recovery of strongylid third stage larvae of parasitic nematodes from small soil samples.

Livestock with access to pasture is generally exposed to infections with parasitic nematode species by uptake of infective third stage larvae (L3) with the grass. L3 can survive on pasture and particularly also in the soil up to several months and sometimes even longer, depending on temperature and humidity. As indicators for health and productivity of grazing animals it is important to determine the intensity and species spectrum of parasitic nematode larvae by analysing grass as well as soil samples. A rapid method for the recovery of L3 using a centrifugal-flotation technique from soil samples of 50-500 g was developed. The method takes advantage of the low specific weight of larvae to separate them from equal sized soil and debris particles by centrifuging them in a saturated sugar solution. A stack of differently sized sieves is used to achieve elimination of larger particles, dust and sugar from the sample to enable easy counting of larvae. Independent of the number of larvae used for inoculation of the samples a mean recovery of 75.3% was obtained. The recovery rates obtained ranged between 60.8% and 88.0% which demonstrates a considerably lower variability compared to earlier approaches and therefore a more precise estimation of the actual numbers of parasite larvae in soil is achieved. Further advantages over already developed methods are the use of easy, affordable and eco-friendly materials, the simplicity of the procedure and a faster processing time with the possibility to examine up to 20 samples per day.

Assessment of the impact of plant species composition and drought stress on survival of strongylid third-stage larvae in a greenhouse experiment.

Grazing livestock is always exposed to infective parasite stages. Depending on the general health status of the animal, the farm management, environmental conditions and pasture exposure, the impact ranges from non-affected to almost moribund animals. The greenhouse experiment was performed to investigate how climatic changes and plant composition influence the occurrence/survival of strongylid third-stage larvae (L3) on pasture. Ten different types of plant species compositions (eight replicates for each) were inoculated with approximately 10,000 Cooperia oncophora L3. The different plant compositions can be assorted to two groups: without legume content and with legume content (52-62% legume content). Half of the replicates were watered adequately, while the other half was hold under drought stress (DS), mimicking longer dry periods. During the DS cycles, the respective containers were not watered until they reached the wilting point. Grass samples were taken 1, 4 and 6 weeks after inoculation, soil samples were taken only once after 6 weeks and all samples were examined for occurrence of L3. After the second DS cycle, the number of L3 present on herbage samples was reduced significantly. The higher the legume content of the pasture composition, the higher is the L3 occurrence on pasture. Independent of the watering scheme, the soil served as the most important reservoir with consistently higher numbers of L3 in the soil compared to herbage.

Suppression of ovine lymphocyte activation by Teladorsagia circumcincta larval excretory-secretory products.

Teladorsagia circumcincta is an important pathogenic nematode of sheep. It has been demonstrated previously that stimulation of murine T lymphocytes with excretory-secretory (ES) products derived from fourth stage larvae of T. circumcincta (Tci-L4-ES) results in de novo expression of Foxp3, a transcription factor intimately involved in regulatory T cell function. In the current study, Foxp3⁺ T cell responses in the abomasum and the effects of Tci-L4-ES on ovine peripheral blood mononuclear cells (PBMC) following T. circumcincta infection were investigated. T. circumcincta infection resulted in a significant increase in numbers of abomasal Foxp3⁺ T cells, but not an increase in the proportion of T cells expressing Foxp3. Unlike in mice, Tci-L4-ES was incapable of inducing T cell Foxp3 expression but instead suppressed mitogen-induced and antigen-specific activation and proliferation of ovine PBMC in vitro. This effect was heat labile, suggesting that it is mediated by protein(s). Suppression was associated with up-regulation of interleukin-10 (IL-10) mRNA, and specific monoclonal antibody neutralisation of IL-10 resulted in a 50% reduction in suppression, indicating involvement of the IL-10 signaling pathway. Suppression was significantly reduced in PBMC isolated from T. circumcincta infected vs. helminth-naïve lambs, and this reduction in suppression was associated with an increase in Tci-L4-ES antigen-specific T cells within the PBMC. In conclusion, we have identified a mechanism by which T. circumcincta may modulate the host adaptive immune response, potentially assisting survival of the parasite within the host. However, the impact of Tci-L4-ES-mediated lymphocyte suppression during T. circumcincta infection remains to be determined.

Phosphoenolpyruvate metabolism in Teladorsagia circumcincta: a critical junction between aerobic and anaerobic metabolism.

Nematodes which have adapted to an anaerobic lifestyle in their adult stages oxidise phosphoenolpyruvate (PEP) to oxaloacetate rather than pyruvate as the final product of glycolysis. This adaptation involves selective expression of the enzyme phosphoenolpyruvate carboxykinase (PEPCK), instead of pyruvate kinase (PK). However, such adaptation is not absolute in aerobic nematode species. We have examined the activity and kinetics of PEPCK and PK in larvae (L(3)) and adults of Teladorsagia circumcincta, a parasite known to exhibit oxygen uptake. Results revealed that PK and PEPCK activity existed in both L(3)s and adults. The enzymes had differing affinity for nucleotide diphosphates: while both can utilise GDP, only PK utilised ADP and only PEPCK utilised IDP. In both life cycle stages, enzymes showed similar affinity for PEP. PK activity was predominant in both stages, although activity of this enzyme was lower in adults. When combined, both the activity levels and the enzyme kinetics showed that pyruvate production is probably favoured in both L(3) and adult stages of T. circumcincta and suggest that metabolism of PEP to oxaloacetate is a minor metabolic pathway in this species.

Recovery of strongylid third-stage larvae from herbage samples: standardisation of a laboratory method and its application in the field.

The estimation of parasitic nematode larvae present on pasture is an important tool applied in many epidemiological studies. In the face of climatic changes, there is increased interest in identifying parameters influencing the survival of free-living stages of parasites under different meteorological conditions. In order to predict possible risk factors for grazing livestock, reliable and reproducible methods to assess the density of larvae on pasture are required. A laboratory method for the recovery of strongylid third-stage larvae from herbage samples was developed, standardised and its efficacy assessed in controlled experiments as well as under field conditions. Grass samples free of any nematode larvae were used and inoculated with known numbers of third-stage larvae of Cooperia oncophora in different concentrations. The grass samples were inoculated with larvae over 24 h, followed by soaking for 4 h. The recovery process included washing over sieves and centrifugation of the obtained liquid. The total time required for the recovery process was about 5-7 h (excluding inoculation). Recovery rates range from 68% to 98% and a strong correlation between numbers of larvae added to the grass samples and numbers of larvae that could be recovered was observed (p < 0.001). The new method proved to be reproducible and provides high recovery rates combined with the potential to handle many samples simultaneously in a relatively short time, thus offering high throughput opportunities applicable to field experiments.

Regional heterogeneity and unexpectedly high abundance of Cooperia punctata in beef cattle at a northern latitude revealed by ITS-2 rDNA nemabiome metabarcoding.

BACKGROUND: The species composition of cattle gastrointestinal nematode (GIN) communities can vary greatly between regions. Despite this, there is remarkably little large-scale surveillance data for cattle GIN species which is due, at least in part, to a lack of scalable diagnostic tools. This lack of regional GIN species-level data represents a major knowledge gap for evidence-based parasite management and assessing the status and impact of factors such as climate change and anthelmintic drug resistance. METHODS: This paper presents a large-scale survey of GIN in beef herds across western Canada using ITS-2 rDNA nemabiome metabarcoding. Individual fecal samples were collected from 6 to 20 randomly selected heifers (n = 1665) from each of 85 herds between September 2016 and February 2017 and 10-25 first season calves (n = 824) from each of 42 herds between November 2016 and February 2017. RESULTS: Gastrointestinal nematode communities in heifers and calves were similar in Alberta and Saskatchewan, with Ostertagia ostertagi and Cooperia oncophora being the predominant GIN species in all herds consistent with previous studies. However, in Manitoba, Cooperia punctata was the predominant species overall and the most abundant GIN species in calves from 4/8 beef herds. CONCLUSIONS: This study revealed a marked regional heterogeneity of GIN species in grazing beef herds in western Canada. The predominance of C. punctata in Manitoba is unexpected, as although this parasite is often the predominant cattle GIN species in more southerly latitudes, it is generally only a minor component of cattle GIN communities in northern temperate regions. We hypothesize that the unexpected predominance of C. punctata at such a northerly latitude represents a range expansion, likely associated with changes in climate, anthelmintic use, management, and/or animal movement. Whatever the cause, these results are of practical concern since C. punctata is more pathogenic than C. oncophora, the Cooperia species that typically predominates in cooler temperate regions. Finally, this study illustrates the value of ITS-2 rDNA nemabiome metabarcoding as a surveillance tool for ruminant GIN parasites.

The vitamin D receptor and inducible nitric oxide synthase associated pathways in acquired resistance to Cooperia oncophora infection in cattle.

Cooperia oncophora is an economically important gastrointestinal nematode in ruminants. Acquired resistance to Cooperia oncophora infection in cattle develops rapidly as a result of prior infections. Naïve cattle, when given a primary infection of high-dose infective L3 larvae, develop a strong immunity to subsequent reinfection. Compared to primary infection, reinfection resulted in a marked reduction in worm establishment. In order to understand molecular mechanisms underlying the development of acquired resistance, we characterized the transcriptomic responses of the bovine small intestine to a primary infection and reinfection. A total of 23 pathways were significantly impacted during infection. The vitamin D receptor activation was strongly induced only during reinfection, suggesting that this pathway may play an important role in the development of acquired resistance via its potential roles in immune regulation and intestinal mucosal integrity maintenance. The expression of inducible nitric oxide synthase (NOS2) was strongly induced during reinfection but not during primary infection. As a result, several canonical pathways associated with NOS2 were impacted. The genes involved in eicosanoid synthesis, including prostaglandin synthase 2 (PTGS2 or COX2), remained largely unchanged during infection. The rapid development of acquired resistance may help explain the lack of relative pathogenicity by Cooperia oncophora infection in cattle. Our findings facilitate the understanding of molecular mechanisms underlying the development of acquired resistance, which could have an important implication in vaccine design.

The dynamics of macroparasite host-self-infection: a study of the patterns and processes of pinworm (Oxyuridae) aggregation.

OBJECTIVES: Among parasites, Taylor's power law identifies a tight relationship in aggregation of macroparasite infection intensity with few exceptions; notably, the nematode family Oxyuridae tends to have higher than expected aggregation. Oxyuridae infect a wide range of mammalian hosts and have a unique reproductive strategy that involves conventional horizontal transmission, as well as re-infection of an already infected host. We asked the question, do the unique aspects of pinworm life-history explain an exception to the widely observed patterns of aggregation of parasite populations? METHODS: We empirically examined the differences among Oxyuridae (genus: Syphacia) compared with other helminth (genus: Heligmosomoides) parasite aggregations in 2 rodent hosts with similar ecology: the yellow-necked mouse (Apodemus flavicollis) from Trento, Italy and the white-footed mouse (Peromyscus leucopus) from Pennsylvania, USA. To investigate the effects of pinworm life-history characteristics on generating aggregation, we present a stochastic model that explores aggregation under a range of host-self-infection, parasite death, and transmission scenarios. RESULTS: Oxyuridae parasites had consistently greater aggregation compared to other nematodes regardless of host or parasite species identity, and pinworm aggregation exceeded the range of macroparasite aggregation described previously. CONCLUSIONS: Our simulations demonstrate that host-self-infection, on its own, is sufficient to generate aggregation values greater than the predicted values.

GTP-cyclohydrolase and development in Teladorsagia circumcincta and Dictyocaulus viviparus (Nematoda: Strongylida).

GTP-Cyclohydrolase (GTP-CH) is necessary for the production of tetrahydrobiopterin, a required cofactor for the three aromatic amino acid hydroxylases and nitric oxide synthases. The gene encoding GTP-CH is transcribed at high levels in infective third larval stages of a number of parasitic trichostrongylid nematodes. We explore the potential role of GTP-CH within the processes of nematode development and environmentally-induced hypobiosis. For two species of parasitic nematode that are of major economic and welfare importance to livestock in temperate regions, Teladorsagia circumcincta and Dictyocaulus viviparus, we have demonstrated that each of the pre-parasitic larval stages transcribe high mean levels of cat-4 (the gene encoding GTP-CH). Using quantitative real-time polymerase chain reaction analysis and two different isolates of D. viviparus, only one of which is capable of entering hypobiosis, we have shown that there were only minor differences between these isolates in mean cat-4 transcript levels, both during the parasitic stages and during the earlier environmental life cycle stages (L(1)-L(3)). Taken together, these data indicate that, although both species of nematode produce high levels of cat-4 transcript in pre-parasitic larval stages, GTP-CH levels are unlikely to be involved in the induction of parasite hypobiosis. Alternative roles for GTP-CH in larval development are discussed.

Development of Graphidium strigosum (Nematoda, Haemonchidae) in its natural host, the rabbit (Oryctolagus cuniculus) and comparison with several Haemonchidae parasites of ruminants.

The morphogenesis (studied for the first time) and the chronology of the life cycle of Graphidium strigosum (Dujardin, 1845) were studied in detail in its natural host, Oryctolagus cuniculus. Naive rabbits were each infected per os with G. strigosum infective larvae (L3). Animals were euthanized each day for the first 10 days after infection (DAI), then every 2 days from 12 to 40 DAI. The free living period lasted 5-8 days at 24°C. By 1 DAI, all the larvae were exsheathed in the stomach. The third molt occurred between 9 and 17 DAI. The last molt occurred between 24 and 32 DAI. The prepatent period lasted 42-44 DAI, while the patent period lasted at least 13 months. For each experiment, the morphology of the different stages of the life cycle was described. The chronology of the G. strigosum life cycle and its morphogenesis were compared to those of different Haemonchidae parasites of ruminants (Ostertagia ostertagi, Teladorsagia circumcincta, Haemonchus contortus, and Haemonchus placei) in their natural hosts.

The comparative morphology of Marshallagia marshalli and Ostertagia occidentalis (Nematoda: Strongylida, Trichostrongylidae) by scanning electron microscopy.

The systematics of the Ostertagiinae is unsettled with no agreement on how many genera and species are present in cattle and sheep. Ten species of Ostertagiinae are commonly parasitic in cattle and sheep. In the global fauna, six of 13 ostertagiine genera are endemic to Iran. The life cycle of Ostertaginae is direct and ingested third-stage larvae after exsheatment in the rumen, penetrate the gastric glands in the abomasal mucosa where two parasitic moults occur before the L5 emerges from the gland. In the present work, Marshallagia marshalli and Ostertagia occidentalis, collected from the abomasums of sheep from Mashhad, Iran, is described. The association of light and scanning electron microscopy (SEM) allowed a detailed analysis of the morphology and ultrastructure of these nematodes. The male body length of M. marshalli and O. occidentalis were 9.3-10.20 and 9.60-10.50 mm, respectively. The female body length of M. marshalli and O. occidentalis were 10.10-15.30 and 10.4-15.70 mm, respectively. One of cervical papillae is seen 333 and 250 µm from the anterior end of male and female body surface in O. occidentalis and 287.5 and 200 µm from the anterior end of male and female body surface in M. marshalli, respectively. The size of cervical papillae is 13.3 µm in male and 10 µm in female in O. occidentalis and 9.33 µm in male and 8.57 µm in female in M. marshalli. Some other taxonomic features of M. marshalli and O. occidentalis, such as details of cephalic region, the system of longitudinal and surface cuticular ridges (synlophe), the orientation of rays of the copulatory bursa, localization of vulva, morphology of vulvar flap, and posterior end of females are also documented by SEM.

Genetic variation among lambs in peripheral IgE activity against the larval stages of Teladorsagia circumcincta.

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.

daf-7-related TGF-beta homologues from Trichostrongyloid nematodes show contrasting life-cycle expression patterns.

The transforming growth factor-beta (TGF-beta) gene family regulates critical processes in animal development, and plays a crucial role in regulating the mammalian immune response. We aimed to identify TGF-beta homologues from 2 laboratory model nematodes (Heligmosomoides polygyrus and Nippostrongylus brasiliensis) and 2 major parasites of ruminant livestock (Haemonchus contortus and Teladorsagia circumcincta). Parasite cDNA was used as a template for gene-specific PCR and RACE. Homologues of the TGH-2 subfamily were isolated, and found to differ in length (301, 152, 349 and 305 amino acids respectively), with variably truncated N-terminal pre-proteins. All contained conserved C-terminal active domains (>85% identical over 115 amino acids) containing 9 cysteine residues, as in C. elegans DAF-7, Brugia malayi TGH-2 and mammalian TGF-beta. Surprisingly, only the H. contortus homologue retained a conventional signal sequence, absent from shorter proteins of other species. RT-PCR assays of transcription showed that in H. contortus and N. brasiliensis expression was maximal in the infective larval stage, and very low in adult worms. In contrast, in H. polygyrus and T. circumcincta, tgh-2 transcription is higher in adults than infective larvae. The molecular evolution of this gene family in parasitic nematodes has diversified the pre-protein and life-cycle expression patterns of TGF-beta homologues while conserving the structure of the active domain.

Teladorsagia circumcincta: survival of adults in vitro is enhanced by the presence of a mammalian cell line.

Adult Teladorsagia circumcincta survival and motility in vitro was examined in a range of different cell culture media, supplements and gas mixes. Under optimum conditions, worms survived for 14 days, exhibiting high motility for 9 days and egg production for 72 h. Optimum conditions involved co-culture of worms with a HeLa cell line in a supplemented cell medium (CEM) and an atmosphere containing 10% CO(2), 5% O(2) 85% N(2), 65% humidity at 37 degrees C. The incubation medium consisted of Minimum Essential Medium with 10% fetal calf serum, 1% non-essential amino acids, 1% glutamax and 1% penicillin-neomycin-streptomycin cocktail mix. Compared with optimum conditions, incubation in CEM alone, cell conditioned CEM, RPMI alone, Medium 199 alone, reduced CO(2) or O(2), or when cells were replaced with Escherichia coli, both survival and motility were reduced. Optimum conditions for adult T. circumcincta maintenance for culture, anthelmintic testing or generation of excretory/secretory products are described.

Long-acting moxidectin for the control of trichostrongylid infections of sheep in south-eastern Australia.

OBJECTIVE: To compare the effectiveness of a single long-acting injection of moxidectin, given in either summer or winter, with a two-summer ('standard') treatment strategy for controlling gastrointestinal nematodes in spring-lambing Merino ewes. DESIGN: A replicated study over 2 years in the high winter rainfall environment of western Victoria. METHODS: Measures of worm infection were worm egg counts of ewes and total worm counts of tracer lambs. Measures of production were body weight, proportion of pregnant ewes and number of weaned lambs. RESULTS: The egg counts of ewes given the standard strategy followed a characteristic pattern, rising to 300 to 600 eggs per gram before the second summer treatment in February. During this time, ewes given long-acting moxidectin in December had zero or negligible counts. There were no consistent differences between tracer worm counts or production measures for the two groups treated in summer, but when data from both years were pooled, the total egg output from December to May was significantly lower for ewes treated with long-acting moxidectin in December. Ewes not treated during summer had lower body weights, and higher worm egg counts and tracer worm counts, showing that this was not a suitable strategy at that time of the year. However, following treatment with long-acting moxidectin in winter, ewes had rapid compensatory weight gain and lower egg output than ewes treated in summer. CONCLUSION: A single injection of long-acting moxidectin in December is an effective treatment strategy for Merino ewes lambing in spring.

Adaptations and phenotypic plasticity in developmental traits of Marshallagia marshalli.

Despite the economic, social and ecological importance of the ostertagiine abomasal nematode Marshallagia marshalli, little is known about its life history traits and its adaptations to cope with environmental extremes. Conserved species-specific traits can act as exaptations that may enhance parasite fitness in changing environments. Using a series of experiments, we revealed several unique adaptations of the free-living stages of M. marshalli that differ from other ostertagiines. Eggs were isolated from the feces of bighorn sheep (Ovis canadensis) from the Canadian Rocky Mountains and were cultured at different temperatures and with different media. Hatching occurred primarily as L1s in an advanced stage of development, morphologically very similar to a L2. When cultured at 20 °C, however, 2.86% of eggs hatched as L3, with this phenomenon being significantly more common at higher temperatures, peaking at 30 °C with 28.95% of eggs hatching as L3s. After hatching, free-living larvae of M. marshalli did not feed nor grow as they matured from L1 to infective L3. These life history traits seem to be adaptations to cope with the extreme environmental conditions that Marshallagia faces across its extensive latitudinal distribution in North America and Eurasia. In order to refine the predictions of parasite dynamics under scenarios of a changing climate, basic life history traits and temperature-dependent phenotypic behaviour should be incorporated into models for parasite biology.