Disease transmission in an extreme environment: nematode parasites infect reindeer during the Arctic winter

Parasitic nematodes are found in almost all wild vertebrate populations but few studies have investigated these host-parasite relationships in the wild. For parasites with free-living stages, the external environment has a major influence on life-history traits, and development and survival is generally low at sub-zero temperatures. For reindeer that inhabit the high Arctic archipelago of Svalbard, parasite transmission is expected to occur in the summer, due to the extreme environmental conditions and the reduced food intake by the host in winter. Here we show experimentally that, contrary to most parasitic nematodes, Marshallagia marshalli of Svalbard reindeer is transmitted during the Arctic winter. Winter transmission was demonstrated by removing parasites in the autumn, using a novel delayed-release anthelmintic bolus, and estimating re-infection rates in reindeer sampled in October, February and April. Larval stages of nematodes were identified using molecular tools, whereas adult stages were identified using microscopy. The abundance of M. marshalli adult worms and L4s increased significantly from October to April, indicating that reindeer were being infected with L3s from the pasture throughout the winter. To our knowledge, this study is the first to experimentally demonstrate over-winter transmission of a gastro-intestinal nematode parasite in a wild animal. Potential mechanisms associated with this unusual transmission strategy are discussed in light of our knowledge of the life-history traits of this parasite.     

Population dynamics of Trichostrongylus colubriformis in sheep: computer model to simulate grazing systems and the evolution of anthelmintic resistance

A computer model was developed to simulate Trichostrongylus colubriformis populations, their level of resistance to the common anthelmintics, host mortalities and acquired immunity. Predictions were based on sheep management practices such as lambing, weaning, sheep/paddock rotation, anthelmintic treatment, the use of controlled release devices (capsules) for anthelmintic delivery and daily meteorological records to determine the development and survival of infective larvae (L3) on pasture. Evolution of drug resistance was determined by a simple genetic system which allowed for up to three genes, each with two alleles, to give a maximum of 27 genotypes associated with one drug or three genotypes for each of three drugs. The model was validated against egg counts, L3 counts on pasture and host mortalities observed in a grazing trial, however, aspects of the model such as the development of drug resistance and use of the model in a variety of climatic zones have yet to be tested against field observations. The model was used to examine the impact of grazing management and capsule use on anthelmintic resistance and sheep production over 20 years using historical weather data. Predictions indicated that grazing management can play a dominant role in parasite control and that capsule use will reduce sheep mortalities and production losses, and in some circumstances will not cause a substantial increase in anthelmintic resistance for up to 5 years.     

Evidence for continued transmission of parasitic nematodes in reindeer during the Arctic winter.

Living in the high Arctic, the Svalbard reindeer (Rangifer tarandus platyrhynchus) and its trichostrongyle nematodes experience a long cold winter from October to late May/early June. Over this period, transmission would be expected to be low. However, in culled reindeer the abundance of infection increased from autumn to late winter, providing evidence for continued transmission within this period. To our knowledge this is the first time this has been demonstrated in a climate with temperatures consistently below 0 degrees C. In one winter (1996-1997), the average fraction of nematodes found as larvae in the abomasal mucosa increased from around 10% to 50% between October and March. This suggests that arrested development took place throughout the winter. We found no evidence for an efficient acquired immune response towards the nematodes. The abundance of infection did not tend to decrease with increasing host age after an earlier peak, but levelled off instead, as predicted by a simple immigration-death model. In the late winter when the nutritional plane is low, both adult reindeer and calves had high worm burdens at intensities that may affect their condition and fitness.     

Leptopilina heterotoma and L. boulardi: strategies to avoid cellular defense responses of Drosophila melanogaster

Eggs of three strains of the cynipid parasitoid Leptopilina heterotoma and a Tunisian strain (G317) of L. boulardi are not encapsulated by hemocytes of Drosophila melanogaster hosts, but the eggs of a Congolese strain (L104) of L. boulardi are encapsulated. To determine the reason for the difference in host response against the parasitoid eggs, lamellocytes (hemocytes that encapsulate foreign objects and form capsules around endogenous tissues in melanotic tumor mutants) were examined in host larvae parasitized by the five Leptopilina strains. Parasitization by the three L. heterotoma strains affected the morphology of host lamellocytes and suppressed endogenous melanotic capsule formation in melanotic tumor hosts. L104 did not alter the morphology of host lamellocytes nor block tumor formation in melanotic tumor mutant hosts. The morphology of some lamellocytes was affected by G317 parasitization but host lamellocytes were still capable of forming melanotic tumors and encapsulating dead supernumerary parasitoid larvae. Therefore, the eggs of strains affecting lamellocyte morphology are protected from encapsulation by the host's blood cells. L. heterotoma eggs float freely in the host hemocoel but L. boulardi eggs are attached to host tissue surfaces. Lamellocytes cannot infiltrate the attachment site so the capsule around the L104 egg remains incomplete. The wasp larva uses this gap in the capsule as an escape hatch for emergence.     

The effect of exogenous testosterone on homospermic and heterospermic fertility in the cock

Cocks were administered testosterone (T) for 14 days via 0, 1, 2, or 4 Silastic capsules implanted subcutaneously. Each capsule released 1.04 mg of T per day. Concentrations of T in plasma and the proportion of eggs fertilized from homospermic insemination of hens were determined. Concentrations of T in plasma were variable and unaffected by treatment. The proportion of eggs that were fertilized by cocks decreased during treatment with 1 capsule, increased over the experiment in the group with 2 capsules, and increased after treatment ended in the group with 4 capsules. In heterospermic tests, cocks with distinguishable offspring were paired and semen was mixed within pairs. One cock in each pair received either 1 or 4 Silastic capsules containing T for 14 days; the other cock in the pair received none. The proportion of chicks sired by cocks treated with 1 capsule remained unchanged throughout the experiment, whereas the proportion sired by cocks treated with 4 capsules decreased markedly during the recovery period. The response to T was apparently dependent upon dosage and the sensitivity of the cock to T. The concentration of T in the plasma of the cock had little relationship to fertility. These results indicate that heterospermic insemination can be used as a sensitive method to detect the subtle effects of hormonal treatment.     

Biology,host preferences and fitness of <Emphasis Type="Italic">Oulema melanopus</Emphasis> (Coleoptera: Chrysomelidae), a recent invasive pest in Western Canada

The developmental parameters and fitness of a recent invasive insect pest of cereals in western Canada, the cereal leaf beetle, Oulema melanopus (L.) (Coleoptera: Chrysomelidae), were compared on live plants and excised leaves of commercial cultivars of potential cereal hosts including wheat (winter and spring), oats, barley, corn, rye and triticale. Host preference and utilization within the fundamental host range of O. melanopus differed. The biological parameters differed significantly when the larvae were reared on excised tissues versus on live host plants. In both studies, wheat (winter and spring), oat (cv. Morgan) and barley were the most preferred hosts in terms of development, survivorship, adult weights and fecundity. Prolonged development with low fitness gains was noted on corn. Although the development on hosts such as rye and triticale was prolonged and adult fitness was low, the survivorship was high on these hosts. Hence, these crops can act as secondary hosts for the beetle in its new eco-region. Larvae fed foliage of Waldern, a local oat cultivar, had increased developmental time and lower survivorship compared with other cereal hosts. Despite differences in fitness gains among hosts, similar numbers of eggs were laid on all hosts. Hence, the oviposition choice of O. melanopus may not be driven by fitness gains alone and it may indicate adaptive strategy to hosts in newer environments.     

Deciphering the role of the capsule of Klebsiella pneumoniae during pathogenesis: A cautionary tale

Extracellular capsule polysaccharides increase the cellular fitness under abiotic stresses and during competition with other bacteria. They are best-known for their role in virulence, particularly in human hosts. Specifically, capsules facilitate tissue invasion by enhancing bacterial evasion from phagocytosis and protect cells from biocidal molecules. Klebsiella pneumoniae is a worrisome nosocomial pathogen with few known virulence factors, but the most important one is its capsule. In this issue, Tan et al. assess the fitness advantage of the capsule by competing a wild-type strain against four different mutants where capsule production is interrupted at different stages of the biosynthetic pathway. Strikingly, not all mutants provide a fitness advantage. They suggest that some mutants have secondary defects altering virulence-associated phenotypes and blurring the role of the capsule in pathogenesis. This study indicates that the K1 capsule in K. pneumoniae is not required for gut colonization but that it is critical for bloodstream dissemination to other organs. These results contribute to clarify the contradictory literature on the role of the Klebsiella capsule during infection. Finally, the varying fitness effects of different capsule mutations observed for K. pneumoniae K1 might apply also to other capsulated diderm bacteria that are facultative or emerging pathogens.     

Rumen microbial diversity in Svalbard reindeer, with particular emphasis on methanogenic archaea

Ruminal methanogens, bacteria and ciliate protozoa of Svalbard reindeer grazing natural pastures in October (late fall) and April (late winter) were investigated using molecular-based approaches. The appetite of the Svalbard reindeer peaks in August (summer) and is at its lowest in March (winter). Microbial numbers, quantified by real-time PCR, did not change significantly between October and April, when food intakes are at similar levels, although the numbers of methanogens tended to be higher in October ( P =0.074), and ciliate numbers tended to be higher in April ( P= 0.055). Similarly, no change was detected in the bacterial and protozoal population composition by rRNA gene-based denaturing gradient gel electrophoresis analysis. Dominant methanogens were identified using a 16S rRNA gene library (97 clones) prepared from pooled PCR products from reindeer on October pasture ( n =5). Eleven of the 22 distinct operational taxonomic units (OTUs) generated exhibited a high degree of sequence similarity to methanogens affiliated with Methanobacteriales (eight OTUs), Methanomicrobiales (one OTU) and Methanosarcinales (two OTUs). The remaining 11 OTUs (53% of the clones) were associated with a cluster of uncultivated ruminal archaea. This study has provided important insights into the rumen microbiome of a high-arctic herbivorous animal living under harsh nutritional conditions, and evidence suggesting that host type affects the population size of ruminal methanogens.     

Male hosts are responsible for the transmission of a trophically transmitted parasite, Pterygodermatites peromysci, to the intermediate host in the absence of sex-biased infection

Field studies have identified that male-biased infection can lead to increased rates of transmission, so we examined the relative importance of host sex on the transmission of a trophically transmitted parasite (Pterygodermatites peromysci) where there is no sex-biased infection. We experimentally reduced infection levels in either male or female white-footed mice (Peromyscus leucopus) on independent trapping grids with an anthelmintic and recorded subsequent infection levels in the intermediate host, the camel cricket (Ceuthophilus pallidipes). We found that anthelmintic treatment significantly reduced the prevalence of infection among crickets in both treatment groups compared with the control, and at a rate proportional to the number of mice de-wormed, indicating prevalence was not affected by the sex of the shedding definitive host. In contrast, parasite abundance in crickets was higher on the grids where females were treated compared with the grids where males were treated. These findings indicate that male hosts contribute disproportionately more infective stages to the environment and may therefore be responsible for the majority of parasite transmission even when there is no discernable sex-biased infection. We also investigated whether variation in nematode length between male and female hosts could account for this male-biased infectivity, but found no evidence to support that hypothesis.     

Release studies of Lactobacillus casei strain Shirota from chitosan-coated alginate-starch microcapsules in ex vivo porcine gastrointestinal contents

AIMS: To investigate the release characteristics of Lactobacillus casei strain Shirota (LCS) from chitosan-coated alginate-starch (CCAS) capsule in different regions of ex vivo porcine gastrointestinal (GI) contents. METHODS AND RESULTS: A 0.1 g of CCAS encapsulated bacteria (containing c. 10(8) CFU of LCS) were incubated in different sections of ex vivo porcine GI contents (10 ml) anaerobically at 37 degrees C. Samples were taken from different GI contents at different time intervals (up to 24 h) and estimated for the release of LCS from capsules. There was almost a complete release (1.1 x 10(8) CFU) of LCS from capsules within 8 h of incubation in ileal contents, while it took nearly 12 h to release the completely encapsulated bacteria in colon content under similar conditions. There was only a partial release of encapsulated LCS, incubated in duodenal and jejunal contents, while there was no significant (P > 0.05) release of encapsulated bacteria in gastric contents even after 24 h of incubation. CONCLUSIONS: The capsules were able to release viable probiotic cells completely in ex vivo porcine ileal and colon contents. SIGNIFICANCE AND IMPACT OF THE STUDY: CCAS capsule can be an effective way of protecting probiotic bacteria from adverse gastric conditions and delivering viable bacteria to the host intestinal systems.     

Seasonal cycles in the output of first stage larvae of the nematode Elaphostrongylus rangiferi from reindeer,Rangifer tarandus tarandus

Summary The output of first stage larvae of the neurotrophic nematode Elaphostrongylus rangiferi was studied both in a herd of reindeer in the field and in reindeer held in captivity. There was a marked seasonal cycle in the output of larvae from infected reindeer. This seasonal cycle is dependent on host sex. After an initial phase of logarithmic increase from the onset of patency in late winter/spring, the larval output declines to a minimum in summer in both female and male reindeer. From then onwards a yearly cycle is repeated with a maximum density of larvae in autumn/early winter from male reindeer, and in late winter/spring from female reindeer. E. rangiferi has an adult longevity of several years in the reindeer, and it is probable that the seasonal cycle of parasite output is linked to seasonal changes in the degree of host stress.     

The developmental strategies and related profitability of an idiobiont ectoparasitoid Sclerodermus pupariae vary with host size

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Dependence of the structure of the capsule surrounding the acanthocephalan Corynosoma strumosum on the species of its natural paratenic host

The fine and ultrafine structure of the capsules surrounding the acanthocephalan Corynosoma strumosum in natural paratenic hosts of three species—the whitespotted greenling Hexagrammos stelleri, Steller’s sculpin Myoxocephalus stelleri, and the saffron cod Eleginus gracilis—was studied. The results of this study, together with earlier data, support the hypothesis that the structure of the capsule depends on the paratenic host species. Three types of capsules differing in the ratio of fibroblasts and inflammatory cells have been identified: fibroblastic (in Japanese smelt, rainbow smelt, and saffron cod), intermediate (in whitespotted greenling), and leukocytic (in Steller’s sculpin and the yellowfin sole). It was assumed that the structure of the capsule is determined by the degree of mutual adaptation of organisms in a given host-parasite system.     

Phylogeography of a parasitoid wasp (Diaeretiella rapae): no evidence of host-associated lineages

The exceptional diversity of insects is often attributed to the effects of specialized relationships between insects and their hosts. Parasite-host interactions are influenced by current natural selection and dispersal, in addition to historical effects that may include past selection, vicariance, and random genetic drift. Both current and historical events can lead to reduced fitness on some hosts. If trade-offs in fitness on alternate hosts are common, adaptation to one host can prevent adaptation to another, giving rise to genetic differentiation among host-associated lineages. Previous studies of Diaeretiella rapae (Hymenoptera: Aphidiidae), a parasitoid of aphids, have revealed additive genetic differences in performance between populations that parasitize different aphid host species. To determine whether D. rapae populations collected from different aphid hosts have diverged into genetically independent lineages, we constructed a haplotype network based on sequence variation in mitochondrial DNA (mtDNA). We used single strand conformation polymorphism (SSCP) analysis to examine 2041 base pairs of mtDNA and to identify nucleotide sequences of 42 unique SSCP haplotypes. We found no association between mtDNA haplotypes and host species in either the ancestral range (Europe, Mediterranean region, Middle East, Asia) or part of the introduced range (western North America). Haplotypes likely to be ancestral were geographically widespread and found on both hosts, suggesting that the ability to use both hosts evolved prior to the diversification of the mtDNA. Ongoing gene flow appears to prevent the formation of host races.     

Functional Anatomy of the Omasum in High Arctic Svalbard Reindeer (Rangifer tarandus platyrhynchus) and Norwegian Reindeer (Rangifer tarandus tarandus)

The structure and fill of the omasum was investigated in summer and in winter in adult female reindeer living on the polar desert and tundra of the high Arctic archipelago of Svalbard and in sub-Arctic mountain habitats in northern Norway The mean total mass of the omasum in non-lactating adult female Svalbard reindeer was 467 g (0.65 g per 100 g live body mass (BM)) in September and 477 g (1.03 g per 100 g BM) in April. By contrast, the mean mass of the omasum in non-lactating adult reindeer in northern Norway was 534 g (0.83 g per 100 g BM) in September but only 205 g (0.35 g per 100 g BM p<0.05) in late March, owing to a decrease in both tissue mass and the wet mass of the contents of the organ. The mean absorptive surface of the omasum in Svalbard reindeer was 2300 cm2 in September and 2023 cm2 in April. In Norwegian reindeer, by contrast, the absorptive surface area decreased from 2201 cm2 in September to 1181 cm2 (p<0.05) in late March. The marked seasonal decline of omasal tissue and contents in Norwegian reindeer probably results from intake of highly digestible forage plants, including lichens, in winter. Svalbard reindeer, a non-migratory sub-species, survive eating poor quality fibrous vascular plants in winter. The absence of any marked seasonal change in the mass, total absorptive surface area or filling of the omasum in Svalbard reindeer in winter despite a substantial decline in body mass presumably reflects their need to maintain maximum absorption of nutrients, including volatile fatty acids, when feeding on such poorly fermentable forage.     

Effects of reindeer on boreal forest floor vegetation: Does grazing cause vegetation state transitions?

Intensive reindeer grazing has been hypothesized to drive vegetation shifts in the arctic tundra from a low-productive lichen dominated state to a more productive moss dominated state. Although the more productive state can potentially host more herbivores, it may still be less suitable as winter grazing grounds for reindeer, if lichens, the most preferred winter forage, are less abundant. Therefore, such a shift towards mosses may have severe consequences for reindeer husbandry if ground-growing lichens have difficulties to recover. We tested if reindeer cause this type of vegetation state shifts in boreal forest floor vegetation, by comparing plant species composition and major soil processes inside and outside of more than 40-year-old exclosures. Lichen biomass was more than twice as high inside exclosures than in grazed controls and almost 5 times higher than in heavily grazed patches. Contrary to our predictions, net N mineralization and plant production were higher in the exclosures than in the grazed controls. The lack of response of phytometer plants in a common garden bioassay indicated that changed soil moisture may drive effects of reindeer on plant productivity in these dry Pine forest ecosystems.     

Host-feeding and oviposition by parasitoids on hosts of different fitness value: Influences of egg load and encounter rate

A dynamic optimization model is presented for the decision to host feed or oviposit on hosts by female parasitoids. Optimal host utilization decisions are compared between two host types with different fitness payoffs for oviposition. The model predicts that hosts of higher fitness value should always be used for oviposition unless the egg load approaches zero. This general prediction is not influenced by levels of host availability or metabolic reserves, the age of the parasitoid, or the magnitude of the fitness difference. An egg-load threshold is predicted above which lower value hosts should be used for oviposition and below which they are used for host feeding. The position of this egg-load threshold is higher when the difference in fitness between host types is larger. The threshold is also higher when overall host availability is high or metabolic reserves for the production of new eggs are low. The threshold for oviposition on low-value hosts decreases to zero near the end of the parasitoid’s life. Under conditions where high-value hosts are rarely encountered compared to low-value hosts, the model predicts that lower value hosts should be accepted for oviposition at a lower egg-load threshold.     

Of mice and worms: are co-infections with unrelated parasite strains more damaging to definitive hosts?

Intraspecific competition between co-infecting parasites can influence the amount of virulence, or damage, they do to their host. Kin selection theory dictates that infections with related parasite individuals should have lower virulence than infections with unrelated individuals, because they benefit from inclusive fitness and increased host longevity. These predictions have been tested in a variety of microparasite systems, and in larval stage macroparasites within intermediate hosts, but the influence of adult macroparasite relatedness on virulence has not been investigated in definitive hosts. This study used the human parasite Schistosoma mansoni to determine whether definitive hosts infected with related parasites experience lower virulence than hosts infected with unrelated parasites, and to compare the results from intermediate host studies in this system. The presence of unrelated parasites in an infection decreased parasite infectivity, the ability of a parasite to infect a definitive host, and total worm establishment in hosts, impacting the less virulent parasite strain more severely. Unrelated parasite co-infections had similar virulence to the more virulent of the two parasite strains. We combine these findings with complementary studies of the intermediate snail host and describe trade-offs in virulence and selection within the life cycle. Damage to the host by the dominant strain was muted by the presence of a competitor in the intermediate host, but was largely unaffected in the definitive host. Our results in this host–parasite system suggest that unrelated infections may select for higher virulence in definitive hosts while selecting for lower virulence in intermediate hosts.     

Winter host exploitation influences fitness traits in a parasitoid

Organisms can either evade winter's unfavourable conditions by migrating or diapausing, or endure them and maintain their activities. When it comes to foraging during winter, a period of scarce resources, there is strong selective pressure on resource exploitation strategy. Generalist parasitoids are particularly affected by this environmental constraint, as their fitness is deeply linked to the profitability of the available hosts. In this study, we considered a cereal aphid–parasitoid system and investigated (1) the host–parasitoid community structure, host availability, and parasitism rate in winter, (2) the influence of host quality in terms of species and instars on the fitness of the aphid parasitoid Aphidius rhopalosiphi De Stefani‐Perez (Hymenoptera: Braconidae: Aphidiinae), and (3) whether there is a detectable impact of host fidelity on parasitism success of this parasitoid species. Host density was low during winter and the aphid community consisted of the species Rhopalosiphum padi L. and Sitobion avenae Fabricius (both Hemiptera: Aphididae), both parasitized by A. rhopalosiphi at non‐negligible rates. Aphidius rhopalosiphi produced more offspring when parasitizing R. padi compared with S. avenae, whereas bigger offspring were produced when parasitizing S. avenae. Although aphid adults and old larvae were significantly larger hosts than young larvae, the latter resulted in higher emergence rates and larger parasitoids. No impact of host fidelity on emergence rates or offspring size was detected. This study provides some evidence that winter A. rhopalosiphi populations are able to take advantage of an array of host types that vary in profitability, indicating that host selectivity may drop under winter's unfavourable conditions.