Developmental instability and immune function in colour polymorphic pygmy grasshoppers

Random left minus right deviations from symmetry in otherwise bilaterally symmetric traits may arise due to developmental instability in response to environmental stress. Here we test for variation in developmental instability, measured as asymmetry of (femur) size, among individuals belonging to four different genetically encoded colour morphs of the pygmy grasshopper Tetrix undulata (Sow.) (Orthoptera: Tetrigidae). Such a difference is expected under the hypotheses that perturbed growth and development and concomitant morphological asymmetry may result from exposure to unfavourable temperatures or costs associated with melanization, or reflect a by-product of past disease in individuals with poor immune responsiveness. Results from mixed model two-way ANOVAs uncovered no statistically significant directional asymmetry in femur size, whereas non-directional asymmetry was significant in each of the four different colour morphs. The degree of intraindividual, interlateral variance in femur size varied significantly among individuals belonging to different colour morphs, being considerably higher in the black morph, which also suffers the greatest risk of overheating, compared to the brown, striped and grey morphs. Asymmetry in femur size was not associated with one measure of immune responsiveness, the encapsulation response of individuals experimentally implanted with a novel antigen consisting of a nylon mono-filament. These results are consistent with the notion that individual pigmentation pattern may indirectly influence developmental instability and morphological asymmetry, via the effects of coloration on body temperature.Co-ordinating editor: Hurst     

Parasite-mediated growth patterns and nutritional constraints in a cavity-nesting bird

1. Trade-offs between growth and immunity of nestling birds can be influenced by parasites, but the magnitude of these effects may depend on availability of critical dietary nutrients. Owing to their importance for both immune system function and growth, dietary carotenoids have the potential to mediate parasite-induced developmental strategies of avian hosts. 2. The effects of ectoparasitic blow flies Protocalliphora spp. and dietary carotenoids (lutein and zeaxanthin) on immune function and patterns of growth in nestling mountain bluebirds Sialia currucoides were investigated by combining parasite removal and carotenoid supplementation treatments in a 2 x 2 design. 3. Supplemental carotenoids enhanced nestlings' T-cell-mediated immune response following intradermal injection of phytohaemagglutinin. 4. The effect of carotenoid supplementation on rate of mass gain depended on whether broods were exposed to parasites: among parasitized broods, those receiving supplemental carotenoids gained mass more rapidly than nonsupplemented broods, whereas there was no effect of supplemental carotenoids on growth of mass in broods that had parasites removed. This suggests that additional dietary carotenoids allowed nestlings to compensate for the otherwise detrimental effects of parasites on mass gain. For length of the eighth primary feather at fledging, early and late broods differed in their response to parasitism: early broods showed an increase in feather length when parasites were removed, while nestlings in late broods had shorter feathers in the absence of parasites. We suggest that this may reflect within-season variation in parasite-mediated growth strategies of nestlings. 5. Maternal condition was positively associated with mass, condition and rate of feather growth of offspring under all conditions, and also influenced nestling immunocompetence, but only in the absence of parasites. 6. We conclude that dietary carotenoids alleviate some of the detrimental effects of parasites on nestling birds; however, parasites also appear to specifically influence other growth and resource allocation strategies, and possibly constrain maternal or genetic effects on offspring phenotype, irrespective of dietary carotenoid availability.     

Equal temperature–size responses of the sexes are widespread within arthropod species

Sexual size dimorphism (SSD) is often affected by environmental conditions, but the effect of temperature on SSD in ectotherms still requires rigorous investigation. We compared the plastic responses of size-at-maturity to temperature between males and females within 85 diverse arthropod species, in which individuals of both sexes were reared through ontogeny under identical conditions with excess food. We find that the sexes show similar relative (proportional) temperature–body size (T–S) responses on average. The high degree of similarity occurs despite an analysis that includes a wide range of animal body sizes, variation in degree of SSD and differences in the sign of the T–S response. We find no support for Rensch''s rule, which predicts greater variation in male size, or indeed the reverse, greater female size variation. SSD shows no systematic temperature dependence in any of the 17 arthropod orders examined, five of which (Diptera, Orthoptera, Lepidoptera, Coleoptera and Calanoida) include more than six thermal responses. We suggest that the same proportional T–S response may generally have equivalent fitness costs and benefits in both sexes. This contrasts with effects of juvenile density, and food quantity/quality, which commonly result in greater size plasticity in females, suggesting these variables have different adaptive effects on SSD.     

Environmental stress increases the prevalence and intensity of blood parasite infection in the common lizard Lacerta vivipara

Parasites affect the life-histories and fitness of their hosts. It has been demonstrated that the ability of the immune system to cope with parasites partly depends on environmental conditions. In particular, stressful conditions have an immunosuppressive effect and may affect disease resistance. The relationship between environmental stress and parasitism was investigated using a blood parasite of the common lizard Lacerta vivipara. In laboratory cages, density and additional stressors had a significant effect on the intensity of both natural parasitaemia and parasitaemia induced by experimental infection. Four weeks after infection, crowded lizards had three times more parasites than noncrowded lizards. After 1 month of stress treatment, naturally infected lizards had a significantly higher level of plasma corticosterone and a higher parasite load than nonstressed individuals. In seminatural enclosures, stress induced by the habitat quality affected both the natural blood parasite prevalence and the intensity of parasitaemia of the host.     

Experimental evidence for latent developmental plasticity: intertidal whelks respond to a native but not an introduced predator

Animals with highly inducible traits may show no inducible response when exposed to a related but wholly novel cue. This appears to be true for the intertidal whelk Nucella lamellosa faced with a voracious introduced predator. In the laboratory, we exposed whelks to effluent from two species of predatory crab, the native red rock crab Cancer productus and the invasive European green crab Carcinus maenas. Nucella and Cancer have a long shared history in the northeast Pacific, whereas potential interaction with Carcinus began here less than 10 years ago. Although Nucella responded adaptively to Cancer effluent by increasing shell thickness and decreasing somatic growth, there was no such response to Carcinus. Furthermore, thicker shelled Nucella were less likely to be eaten by Carcinus. Because Nucella produces thicker shells when exposed to Cancer cues, its ability to respond similarly to Carcinus depends only on the coupling of the Carcinus cue to the existing developmental pathways for adaptive changes in shell form. Such coupling of latent plasticity to a novel cue -- via genetic changes or associative learning -- could explain many cases of rapid phenotypic change following a sudden shift in the environment.     

Interaction between Oesophagostomum columbianum and Oesophagostomum venulosum in sheep

Interactions between Oesophagostomum columbianum and Oesophagostomum venulosum were studied by comparing the establishment, development and distribution of each species in single and mixed infections in sheep. The establishment of O. venulosum was reduced by 90 % in sheep previously infected with O. columbianum and by a mean of 50 % in sheep given a simultaneous concurrent infection with O. columbianum. Prior or simultaneous infection with O. venulosum had little or no effect on the establishment of O. columbianum. It is concluded that the increase in incidence of O. venulosum in sheep on the Northern Tablelands of New South Wales has been consequent upon but not the cause of the decline in incidence of O. columbianum. The decline in O. columbianum has probably occurred because of a combination of factors including the failure of infective larvae to overwinter on pasture, the use of efficient anthelmintics and changes in pasture composition.     

Effects of food availability on predator-induced morphological defence in the ciliate Euplotes octocarinatus (Protista)

The effect of a prey's food resources on predator-induced morphological transformation was studied in two ciliate species: Euplotes octocarinatus (prey) and Stylonychia mytilus (predator). The ability to produce defence in Euplotes was much reduced by prolonged starvation. The extent of prey morphological transformation and therefore the effectiveness of the defence was inversely related to the length of time without food. The results suggest that the defence involves an energetic cost which may be a significant part of the whole cell budget, at least when food is in short supply. Not only the morphological transformation but also the maintenance of the anti-predator phenotype were energetically costly for Euplotes, the cost being proportional to the defence level. The morphological transformation was affected by the kind of food (algae) provided to the prey. Ciliates fed Chlorogonium elongatum attained their maximum width sooner than those fed Chlamydomonas. Thus, both the quantity and the quality of the food available may significantly affect the magnitude of response and therefore the potential success of an induced defence.     

The effect of predator and prey density on the induced defence of a ciliate

1. The level of antipredator defence should be proportional to the actual attack probability to minimize the cost of defence and maximize the net benefit.
2. The hypothesis that the induced antipredator morphology of Euplotes octocarinatus is a graded response to the actual risk of predation by Stylonychia mytilus was tested by manipulating the density of both prey and predator populations.
3. The magnitude of the response was graded according to both predator and prey density. A dense prey population may be protective since a prey is more exposed to a predator's attack as a solitary individual.
4. The results suggest that Euplotes is able to 'estimate' the real risk of predation and respond appropriately, without mobilizing more resources than needed.
5. Separation of the prey and predator with a nylon net revealed that the response was not induced by a water-transmitted factor but that direct cell-to-cell contacts were important. This finding departs from those of other studies.     

Multiple introductions of the Spiroplasma bacterial endosymbiont into Drosophila

Bacterial endosymbionts are common in insects and can have dramatic effects on their host's evolution. So far, the only heritable symbionts found in Drosophila have been Wolbachia and Spiroplasma . While the incidence and effects of Wolbachia have been studied extensively, the prevalence and significance of Spiroplasma infections in Drosophila are less clear. These small, gram-positive, helical bacteria infect a diverse array of plant and arthropod hosts, conferring a variety of fitness effects. Male-killing Spiroplasma are known from certain Drosophila species; however, in others, Spiroplasma appear not to affect sex ratio. Previous studies have identified different Spiroplasma haplotypes in Drosophila populations, although no extensive surveys have yet been reported. We used a multilocus sequence analysis to reconstruct a robust Spiroplasma endosymbiont phylogeny, assess genetic diversity, and look for evidence of recombination. Six loci were sequenced from over 65 Spiroplasma -infected individuals from nine different Drosophila species. Analysis of these sequences reveals at least five separate introductions of four phylogenetically distinct Spiroplasma haplotypes, indicating that more extensive sampling will likely reveal an even greater Spiroplasma endosymbiont diversity. Patterns of variation in Drosophila mitochondrial haplotypes in Spiroplasma -infected and uninfected flies imply imperfect vertical transmission in host populations and possible horizontal transmission.     

Collective defence portfolios of ant hosts shift with social parasite pressure

Host defences become increasingly costly as parasites breach successive lines of defence. Because selection favours hosts that successfully resist parasitism at the lowest possible cost, escalating coevolutionary arms races are likely to drive host defence portfolios towards ever more expensive strategies. We investigated the interplay between host defence portfolios and social parasite pressure by comparing 17 populations of two Temnothorax ant species. When successful, collective aggression not only prevents parasitation but also spares host colonies the cost of searching for and moving to a new nest site. However, once parasites breach the host''s nest defence, host colonies should resort to flight as the more beneficial resistance strategy. We show that under low parasite pressure, host colonies more likely responded to an intruding Protomognathus americanus slavemaker with collective aggression, which prevented the slavemaker from escaping and potentially recruiting nest-mates. However, as parasite pressure increased, ant colonies of both host species became more likely to flee rather than to fight. We conclude that host defence portfolios shift consistently with social parasite pressure, which is in accordance with the degeneration of frontline defences and the evolution of subsequent anti-parasite strategies often invoked in hosts of brood parasites.