Per Johan Jakobsen

Male-biased susceptibility to helminth infection : an experimental test with a copepod

Recent reviews on sex biases in parasitism found that males generally suffer from a slightly higher susceptibility to helminth infection than females. Sex hormones like testosterone have been suggested as the mechanism that causes the bias in parasite susceptibility. Many of the original studies used data that stem from free-ranging host populations which were naturally infected. Since gender-related behaviours could result in differential exposure to parasites, it is necessary to experimentally disentangle differences in exposure from differences in susceptibility. We tested whether we could find a sex bias in susceptibility under experimental conditions, i.e. by controlling for gender related differences in exposure. Furthermore, we used an invertebrate host to test whether the pattern observed in many vertebrates can be extended to a host species that lacks testosterone. As models we used the cestode Schistocephalus solidus and its first intermediate host, the copepod Macrocyclops albidus. We found that male copepods were more often infected than females. Since infected males had on average also more parasite larvae than infected females, the chance of a S. solidus larvae to become established in its first intermediate host was more than twice as high in male than in female copepods. This shows that, under experimental conditions, male-biased susceptibility to helminth infection can be very strong. Moreover, the pattern seen in many vertebrates can be extended to an invertebrate host that lacks testosterone.

Salmon lice, Lepeophtheirus salmonis, infestation as a causal agent of premature return to rivers and estuaries by sea trout, Salmo trutta, juveniles

A field experiment conducted in the River Lønningdalselven in spring 1992 supports the hypothesis that salmon lice, Lepeophtheirus salmonis, infestations may cause premature return of sea trout, Salmo trutta, juveniles, either to estuaries or to rivers. When lice infested (exposed) and uninfested (control) sea trout juveniles (post smolts) were released simultaneously into the sea, exposed fish returned to the estuarine area earlier compared with controls. Within the following two days, exposed sea trout migrated further into freshwater. At that time they were infested with a median of 62.5 lice, dominated by chalimus larvae and late juveniles. Exposed sea trout suffered from an osmoregulatory failure in sea water and this is considered one reason for infested fish returning to brackish water. While only a few control fish returned to the estuary on the day of release, some more returned to freshwater the following four days. During this time they had become heavily infested with copepodids, and carried a median of 150.0 lice. It is suggested that physiological stress and high infection pressure in the sea results in sea trout juveniles returning to estuaries and freshwater.

The influence of food limitation on swarming behaviour in the waterflea Bosmina longispina

Abstract Bosmina longispina showed a high degree of overdispersal in nature, which drastically reduced the local amount of food within the swarms. In experiments, swarming behaviour occurred only in the light and when food was abundant. In darkness and when food was limiting, B. longispina were more evenly distributed. If swarming prey individuals benefit from a reduced predation risk, the swarming behaviour of B. longispina can be interpreted as an adaptive diurnal rhythm in the trade-off between reducing risk of predation and maximizing feeding rate. Maximal feeding seems to have priority over passive avoidance mechanisms such as swarming when the risk of starvation is high.

Size-specific protection against predation by fish in swarming waterfleas, Bosmina longispina

Abstract When feeding on non-swarming waterflea prey, sticklebacks, Gasterosteus aculeatus, selected larger items. However, when feeding on swarming prey, the fish fed mainly at the edges of the swarms and smaller items were consumed. This selection of small prey items can be explained by the fact that when attacked by a stickleback the waterfleas escaped towards the centre of the swarm, larger individuals more quickly than smaller ones, thus leaving the stickleback with smaller than average prey at the edge of the swarm.

Behavioural response of the water flea Daphnia pulex to a gradient in food concentration

Abstract Individual Daphnia pulex were allowed to distribute themselves in two connected flow-through chambers, only one of which received an input of food. The amount of inflowing food was regulated to give either a high or a low concentration. Daphnia pulex offered high food concentrations were randomly distributed between the two chambers, in spite of differences in concentration of food. However, when food concentrations were low and limiting, the animals responded to the food levels and distributed themselves to a larger extent in the food-enriched chamber.

Fish kairomone regulation of internal swarm structure in Daphnia pulex (Cladocera: Crustacea)

In this laboratory experiment, swarms of D. pulex were artificially created by placing thirty individuals in small chambers containing 130 ml of water. The swimming behaviour of the animals was measured using a video camera and a programme to digitize observations. The D. pulex swimming in water with fish kairomones had a more uniform swimming speed compared with animals in control water. If aggregated prey individuals benefit from a reduced predation risk, and this risk is further reduced by uniformity of swarm members, the uniformity of swimming speed can be interpreted as an behavioural adjustment to minimize the vulnerability to predation.

Effects of the Emotion System on Adaptive Behavior

A central simplifying assumption in evolutionary behavioral ecology has been that optimal behavior is unaffected by genetic or proximate constraints. Observations and experiments show otherwise, so that attention to decision architecture and mechanisms is needed. In psychology, the proximate constraints on decision making and the processes from perception to behavior are collectively described as the emotion system. We specify a model of the emotion system in fish that includes sensory input, neuronal computation, developmental modulation, and a global organismic state and restricts attention during decision making for behavioral outcomes. The model further includes food competition, safety in numbers, and a fluctuating environment. We find that emergent strategies in evolved populations include common emotional appraisal of sensory input related to fear and hunger and also include frequency-dependent rules for behavioral responses. Focused attention is at times more important than spatial behavior for growth and survival. Spatial segregation of the population is driven by personality differences. By coupling proximate and immediate influences on behavior with ultimate fitness consequences through the emotion system, this approach contributes to a unified perspective on the phenotype, by integrating effects of the environment, genetics, development, physiology, behavior, life history, and evolution.

Reciprocal cross infection of sticklebacks with the diphyllobothriidean cestode Schistocephalus solidus reveals consistent population differences in parasite growth and host resistance

BackgroundIn host-parasite evolutionary arms races, parasites are generally expected to adapt more rapidly, due to their large population sizes and short generation times. There exist systems, though, where parasites cannot outpace their hosts because of similar generation times in both antagonists. In those cases concomitant adaptation is expected.MethodsWe tested this hypothesis in the three-spined stickleback-Schistocephalus solidus tapeworm system, where generation times are comparable in both organisms. We chose two populations of sticklebacks which differ prominently in the prevalence of S. solidus and consequently in its level of selective pressure. We performed a full factorial common garden experiment. Particularly, Norwegian (NO) and German (DE) sticklebacks, as well as hybrids between both stickleback populations and in both parental combinations, were exposed each to a single S. solidus originating from the same two host populations.ResultsWe found the infection phenotype to depend on the host population, the parasite population, but not their interaction. NO-parasites showed higher infectivity than DE-parasites, with NO-sticklebacks also being more resistant to DE-parasites than to the sympatric NO-parasite. Reciprocally, DE-hosts were more susceptible to the allopatric NO-parasite while DE-parasites grew less than NO-parasites in all stickleback groups. Despite this asymmetry, the ratio of worm to host weight, an indicator of parasite virulence, was identical in both sympatric combinations, suggesting an optimal virulence as a common outcome of parallel coevolved systems. In hybrid sticklebacks, intermediate infection rates and growth of S. solidus from either origin suggests a simple genetic basis of resistance. However, comparison of infection phenotypes in NO-maternal and DE-maternal hybrid sticklebacks indicates local adaptation to the sympatric counterpart in both the host and the parasite.ConclusionsHost-parasite systems with similar generation time show evidence for concomitant reciprocal adaptation resulting in parasite optimal virulence and host parasite specific resistance.

The emotion system promotes diversity and evolvability

Studies on the relationship between the optimal phenotype and its environment have had limited focus on genotype-to-phenotype pathways and their evolutionary consequences. Here, we study how multi-layered trait architecture and its associated constraints prescribe diversity. Using an idealized model of the emotion system in fish, we find that trait architecture yields genetic and phenotypic diversity even in absence of frequency-dependent selection or environmental variation. That is, for a given environment, phenotype frequency distributions are predictable while gene pools are not. The conservation of phenotypic traits among these genetically different populations is due to the multi-layered trait architecture, in which one adaptation at a higher architectural level can be achieved by several different adaptations at a lower level. Our results emphasize the role of convergent evolution and the organismal level of selection. While trait architecture makes individuals more constrained than what has been assumed in optimization theory, the resulting populations are genetically more diverse and adaptable. The emotion system in animals may thus have evolved by natural selection because it simultaneously enhances three important functions, the behavioural robustness of individuals, the evolvability of gene pools and the rate of evolutionary innovation at several architectural levels.

In vitro transition of Schistocephalus solidus (Cestoda) from coracidium to procercoid and from procercoid to plerocercoid

With the present study, a culture system for successive life-cycle stages of the tapeworm Schistocephalus solidus was developed and this report documents for the first time, cultivation of the procercoid stage of S. solidus from eggs. Additionally we have transformed procercoids dissected from experimentally infected copepods and cultured procercoids into the early plerocercoid stage in vitro. Observations in the culture suggest that the coracidia can interact with their external environment and need no host specific stimuli, except for the components in the culture medium, for activation and hatching from the embryophore. Increasing the culture medium pH from 7.3 to 8.0 improved escape rates and frequencies of hook contractions, suggesting that the oncosphere may recognize and respond to environmental conditions along the host intestine. Procercoids in the culture did not stop growing indicating that conditions within the copepod may be important to limit growth and to induce transformation to plerocercoids. When procercoids are dissected from copepods and transferred to the culture, the outer tegument layers and cercomer starts to loosen. Comparison of the lectin staining of the loosened outer tegument layers and cercomer in procercoids dissected from copepods confirms that transitions of both, the oncosphere to procercoid and procercoid to plerocercoids, has taken place in the in vitro cultures.