Coevolution of parasite virulence and host life history

Most models about the evolutionary interactions between a parasite's virulence and its host's life history neglect two potentially important aspects: epidemiological and coevolutionary feedback. We emphasize their importance by presenting models that describe the coevolution of a semelparous host's age at reproduction and a parasite's virulence in different environmental conditions. In particular, we first show that an epidemiological feedback will lead to a nonmonotonic response of the host's age at reproduction as virulence increases. We then show that the coevolutionary pressure on virulence can lead to complex associations between the host's life history and the parasite's virulence, which would not be expected with more traditional models of host or parasite evolution. Thus, for example, a high mortality rate of the host favours avirulent parasites and late reproduction of the host when the environmental conditions allow the host to grow rapidly, but early reproduction and high virulence when growth is slow.     

Dietary effects on life history traits in a terrestrial isopod: the importance of evaluating maternal effects and trade-offs

Studies of life history aim to explain patterns in the evolution of reproductive investment, growth, and survival. Trade-offs between traits are a fundamental component of life history theory. In herbivorous arthropods life history traits are often responsive to variation in numerous environmental factors, especially diet quality. Using three artificial diets under controlled laboratory conditions, we examined changes in life history traits (i.e. growth rate, offspring number, offspring size, incubation period), trade-offs between traits, and maternal effect on the growth rate of offspring, in the common woodlouse (terrestrial isopod), Porcellio laevis. The high protein diet had significant impacts on offspring production, triggering a smaller-sized offspring, and demonstrating a trade-off between these last two traits. The high carbohydrate diet seldom exerted a significant effect on incubation period. The quality of dietary items evidently has important consequences on the life history of the mother and, thus, on offspring growth; the directions of these effects, however, were opposite. Mothers fed diets with high protein concentrations presented significant maternal effects, measured as offspring growth rate during later ontogeny. Our results support the notion that protein, rather than carbohydrate, concentrations in the diet limit herbivorous arthropods, and have significant consequences on life history traits, as was seen for P. laevis. Clearly, the change in phenotypic correlations between incubation period and offspring number from negative to positive is an empirical demonstration of the context dependence of life history trait trade-offs.     

On the development and reproduction of Botryllus schlossen (Tunicata) colonies from the eastern Mediterranean Sea: plasticity of life history traits

Summary

Analysis of developmental patterns of Botryllus schlosseri colonies from the eastern Mediterranean coast has been performed on 143 genets under different temperature regimens (15,20,27 °C) for up to 22 weeks. While the average maximal size as the length of the blastogenic cycle (BC) varied in respect to water temperature, ontogeny at all temperatures was characterized by 4 developmental stages: 1. the lag phase (the first 1–3 BCs, 1 bud/BC), 2. the exponential growth phase (5.5–7.8 BCs, up to 3 buds/BC), 3. the plateau phase, (7.4–8.2 BCs, 1 bud/BC), 4. the degenerative or the variable phase. Many (40.6%) colonies were not sexually reproductive, the others were male only (30.1%) or hermaphrodites. Colonies at the peak of reproduction develop 1.2 oocytes/zooid, and up to 57.7% produced >4 clutches. Analysing onset of reproduction with maximal colony size revealed 4 patterns, two for “male only” colonies and two for hermaphrodites that varied at different sea water temperatures. In two patterns, the onset of reproduction precedes colony maximal size, and in the others it starts at the peak size or thereafter. Zooids at the colony's periphery developed almost twice as many buds as did zooids at the center, but produced significantly lower numbers of eggs. Fragmentation was recorded in large colonies and was temperature dependent. Results are compared with the data available on populations from other localities indicating for dramatically different developmental modes characteristic to this cosmopolitan species.     

The life history of Sulcascaris sulcata (Nematoda: Ascaridoidea), a parasite of marine molluscs and turtles

Berry G. N. and Cannon L. R. G. 1981. The life history of Sulcascaris sulcata (Nematoda: Ascaridoidea), a parasite of marine molluscs and turtles. International Journal for Parasitoiogy11: 43–54. The morphology, development and hatching of Sulcascaris sulcata eggs are described. Two moults occurred in the egg. Third stage larvae spontaneously hatched and were found to develop in marine bivalves and gastropods. Larvae grew steadily and after three to four months, when about 5 mm long, they moulted to fourth stage larvae characteristic of natural infections in bivalves from commercial catches. Experimentally, when fed to laboratory-reared Caretta caretta, the fourth stage larvae first attached at the oesophago-gastric junction where they moulted to adults in 7–21 days. Subsequent growth to mature adults was obtained by at least 5 months after infection. It is suggested that under natural conditions the life history may take up to 2 years to complete. These findings are discussed in relation to the predatory mode of feeding and the breeding habits of C. caretta and the significance of a possible health hazard to man.     

Dose and host characteristics influence virulence of ranavirus infections

Parasites play a prominent role in the ecology, evolution, and more recently, conservation of many organisms. For example, emerging infectious diseases, including a group of lethal ranaviruses, are associated with the declines and extinctions of amphibians around the world. An increasingly important basic and applied question is: what controls parasite virulence? We used a dose-response experiment with three laboratory-bred clutches of tiger salamander larvae (Ambystoma tigrinum) to test how the size of inoculum and host genetic factors influence the dynamics and outcome of ranavirus infections. We found that infection rates increased with dose and were strongly affected by clutch identity and host life history stage. Case mortality increased with dose of inoculum, but was unaffected by host characteristics. Average survival time decreased with dose and differed among clutches, but this was largely due to differences in the time to onset of symptoms. Overall, our results suggest that dose of inoculum and host characteristics (life history stage and genetic background) influence the establishment and early virus replication, and therefore the virulence of ranavirus infections.     

Host diet as a determinant of parasite growth, reproduction and survival

Changes have been observed in the biology of protozoan and helminth parasites of small mammals during a variety of experimental nutritional manipulations ranging from deficiency in a specific nutrient to the general withholding of food from the host. Commonly observed effects include retardations in parasite growth rate, gametogenesis, fecundity, and asexual multiplication. The duration of patency and of the association of a parasite with its host have also been observed to be curtailed. The nutritional responses of parasites during host hibernation require investigation and much research will be needed to explain how perturbations in host dietary composition and intake may lead to the observed effects. Care should be exercised over applying the results from experimental studies to naturally occurring infections, but the experiments are justified because they indicate potential effects, if not their biological significance, and because causation can be determined.     

Change across the lifespan in a psychological measure of life history strategy

Life history theory predicts that people calibrate their reproductive strategies to local levels of environmental harshness and unpredictability. While previous research has established the importance of early life cues in the development of life history strategy, the degree to which life history strategy exhibits plasticity later in life is unclear. Using longitudinal data (total N = 479) from four archival studies and a recently validated psychological measure of life history strategy, we examined mean-level trends in life history strategy at the level of psychological phenotype between the ages of 7 and 60 and found that life history strategy slowed down linearly as a function of age. Highlighting the importance of sexual selection in shaping life history strategy, we also found that men had a faster life history strategy than women at all ages and that the magnitude of this difference was constant across the lifespan. Our findings suggest that life history strategy development continues even in older adulthood. We discuss the possibility that this occurs in response to the accumulation of biological and social (e.g. offspring, relationships) capital and information about local risks and incentives.     

The effects of background mortality on optimal reproduction in a seasonal environment

We consider optimal annual routines of reproductive behaviour in a seasonal environment. In our model the condition of the organism is adversely affected by hard work, but can recover during easy periods. Our analysis concentrates on the effects of background mortality (i.e., mortality that cannot be avoided) on the optimal strategy and how often an organism following this strategy breeds. In particular, we are concerned with whether reproduction occurs at specific times of year (entrained to the annual cycle), and if so then how many reproductive bouts occur per year. We find that an increase in background mortality can have various effects. If the animal is entrained to the annual cycle and has one breeding attempt per year, then breeding tends to occur earlier and there may be two breeding attempts per season. Another possible outcome is that breeding is no longer entrained. If the animal is entrained but sometimes skips reproduction so that it does not breed every year, then an increase in mortality may make it more likely that the animal breeds every year. We show that as background mortality increases the resultant increase in the frequency of breeding contributes to the increase in annual mortality. We also explore the effects of mortality on the timing of reproduction within a year, highlighting the tension between the interests of the parent and that of the young.     

Ultrastructural study on the spermiogenesis and spermatozoon of the metacercariae of Microphallus primas (Digenea), a parasite of Carcinus maenas

The thread-like spermatozoon of the crab parasite Microphallus primas was studied by electron microscopy. A survey of the head region of the spermatozoon reveals three features hitherto unknown in Platyhelminthes spermatozoa. The first is the aberrant inclusion of the nucleus within one of the two axonemes, limited to the head end region. The second is the coexistence, in the same axoneme, of two patterns, 9 + 0 (doublets without dynein arms) and 9 + "1". The third is the presence of a layer of cortical microtubules running longitudinally from the zone where the nucleus goes from axoneme to the tail region (where the two flagella start). The sequence of events in spermatogenesis is similar to that described for most of the Digenea trematodes, and the spermiogenesis process conforms to a common plan in nearly the whole group.     

The impact of diapause on the evolution of other life history traits in flesh flies

Summary Flesh flies (Sarcophagidae) collected in Costa Rica and Panama lack the pupal diapause that is characteristic of flesh flies from the temperate zone and tropical Africa. The absence of a diapause capacity in the neotropical species correlates with several other life history traits: in most species the post feeding wandering phase of the third larval instar is longer and duration is more variable, adult life is longer, clutch size is smaller, and more clutches are produced. Among species that have the capacity for diapause, risk is invested primarily in the diapausing stage and other life stages are brief. Though diapausing species are short-lived, they produce as many or more progeny than nondiapausing species by increasing clutch size. The slower and more variable developmental rate and increased adult longevity desynchronizes development and permits the nondiapausing species to spread an environmental risk over different stages of the life cycle, thus offering an alternative to diapause. Other traits such as body size, developmental velocity, thermal constant thresholds, thermal constants, age at first reproduction, and the interval between clutches do not appear related to the capacity for diapause.     

Complex and interactive effects of ocean acidification and warming on the life span of a marine trematode parasite

Human activities have caused an increase in atmospheric CO₂ over the last 250 years, leading to unprecedented rates of change in seawater pH and temperature. These global scale processes are now commonly referred to as ocean acidification and warming, and have the potential to substantially alter the physiological performance of many marine organisms. It is vital that the effects of ocean acidification and warming on marine organisms are explored so that we can predict how marine communities may change in future. In particular, the effect of ocean acidification and warming on host-parasite dynamics is poorly understood, despite the ecological importance of these relationships. Here, we explore the response of one himasthlid trematode, Himasthla sp., an abundant and broadly distributed species of marine parasite, to combinations of elevated temperature and pCO₂ that represent physiological extremes, pre-industrial conditions, and end of century predictions. Specifically, we quantified the life span of the free-living cercarial stage under elevated temperature and pCO₂, focussing our research on functional life span (the time cercariae spend actively swimming) and absolute life span (the period before death). We found that the effects of temperature and pCO₂ were complex and interactive. Overall, increased temperature negatively affected functional and absolute life span, e.g. across all pCO₂ treatments the average time to 50% cessation of active swimming was approximately 8 h at 5 °C, 6 h at 15 °C, 4 h at 25 °C, and 2 h at 40 °C. The effect of pCO₂, which significantly affected absolute life span, was highly variable across temperature treatments. These results strongly suggest that ocean acidification and warming may alter the transmission success of trematode cercariae, and potentially reduce the input of cercariae to marine zooplankton. Either outcome could substantially alter the community structure of coastal marine systems.     

Differences in the effects of parental age on offspring life history between tropical and temperate populations of milkweed bugs (Oncopeltus spp.)

Summary We compare the effects of parental age on several offspring life history traits in two milkweed bug populations: a typically univoltine population ofOncopeltus fasciatus from the Sacramento Valley of California and a typically multivoltine population ofO. cingulifer from Monteverde, Costa Rica. Reared under identical conditions (27°C, 12 h light: 12 h dark photoperiod), these bugs exhibit significant differences in the effects of parental age on offspring life history. As they age,O. fasciatus females from Sacramento lay clutches of eggs of decreasing average weight and a decreasing proportion of their eggs produce offspring that survive to adulthood. Those offspring which do survive have a significantly faster developmental rate and females have a larger body size at adult eclosion. AsO. cingulifer from Monteverde age, they also produce lighter eggs, but there is no significant change in the offspring developmental rate, survival or female adult body size. We suggest that these results are largely explicable as the consequence of different parental investment strategies associated with the predictable relationship between parental age and time of season inO. fasciatus but not inO. cingulifer. AsO. fasciatus from Sacramento age, they may be increasing their investment per developing offspring so as to increase the probability that nymphs hatching late in the season will reach a prereproductive adult diapause before the first killing frost.     

Life history evolution in Asobara tabida: plasticity in allocation of fat reserves to survival and reproduction

Life history variation can be genetically based, or it may be due to environmental effects on the phenotype. In this paper we examine life history variation in relation to differences in habitat in the parasitoid Asobara tabida. Differences in the spatial distribution of host patches, the length of the season, host suitability, and competition between parasitoids all contribute to the selection for differences in life history between strains from northern and southern Europe. Strains were compared with respect to the allocation between reproduction and survival in experimental environments that varied with regard to the availability of food or to the number of hosts offered per day. Upon emergence parasitoids originating from southern Europe have higher egg loads and smaller fat reserves than parasitoids originating from northern Europe. Parasitoids from both southern and northern Europe show plasticity in allocation: food shortage causes a decrease in egg production, and rich breeding opportunities an increase in egg production. This plasticity is greater in the northern strain. Fat reserves play a central role in the physiology of this allocation. Fat can be used for both reproduction and survival. There is no oosorption, so once fat is allocated to reproduction it can no longer be used for survival. Differences in plasticity in allocation can therefore be considered as differences in the timing of egg production.     

Host mortality, predation and the evolution of parasite virulence

One of the most accepted views in the theoretical literature on virulence evolution is that a parasite's virulence will evolve to higher levels when its host's background mortality rate increases. Surprisingly, however, although many sources of background mortality involve predation, there has not yet been any theoretical research that explicitly considers how the dynamics of this important ecological interaction affects virulence evolution. Here, we consider how predation affects virulence evolution by explicitly introducing a predator into a classical susceptible–infected–susceptible epidemiological model. We find that, contrary to previous predictions, different sources of host mortality affect virulence evolution in different ways. Moreover, the way in which virulence evolution is affected depends on how tightly coupled the predator's dynamics are to the host population, and this can result in somewhat counterintuitive results. For example, indirect ecological effects can cause elevated host mortality to result in the evolution of lower parasite virulence, even if this elevated mortality arises from factors unrelated to predation.     

Developmental response to a seasonal time constraint: the effects of photoperiod on reproduction in the grasshopper Romalea microptera

Abstract.  1. Some organisms respond adaptively to seasonal time constraints by altering development time to life-history transitions (e.g. metamorphosis, oviposition). Such life-history changes may have costs (e.g. reduced fecundity, mass, offspring quality).
2. The hypothesis that a northern population of the grasshopper Romalea microptera (Beauvois) would show adaptive plasticity in oviposition timing in response to seasonal time constraints was tested by manipulating photoperiod to simulate the middle of the active season (Long photoperiod), the end of the active season (Short photoperiod), and seasonal change (photoperiod Declining from long to short). Females received either high or low food rations. Short or Declining photoperiod were predicted to induce early oviposition with costs of reduced egg number, post-oviposition mass, or egg size, particularly in low-food females.
3. Effects of food ration and photoperiod, but not interaction, were significant in failure time analysis of age at oviposition. mancova on age at oviposition, egg number, and post-oviposition mass yielded similar effects. The multivariate effect of photoperiod resulted primarily from reduced time to oviposition in Short or Declining photoperiod. No costs in egg number or post-oviposition mass were associated with this photoperiod-induced reduction in time to oviposition. The multivariate effect of food ration resulted mainly from lower egg number with low food. Food ration affected egg size, but photoperiod and interaction did not. In all cases, Short and Declining photoperiod produced similar effects.
4. In its northern range, R. microptera accelerates reproduction in response to seasonal constraints, a response that may be adaptive. How R. microptera avoids costs associated with this reduced pre-oviposition period remains unknown.     

Life history strategies and psychopathology: the faster the life strategies,the more symptoms of psychopathology

There is little extant empirical literature examining the associations between life history strategies and symptoms of psychopathology. The current study (N = 138) investigated the associations between life history strategies, symptoms of psychopathology, aggression, incidence of self-harm behaviour, and attachment (perceived parental support) in sample drawn from the general population and community mental health service providers. The results from the study indicate those with a faster life strategy report greater levels of aggression and symptoms of psychopathology. Further, perceptions of poorer parental support were associated with a faster life history strategy. Implications for life history theory, conceptualising psychopathology, and future research directions are discussed.     

The effects of water availability on the life history of the desert snail,Trochoidea seetzeni

Summary In the Negev desert of Israel, the pulmonate land snailTrochoidea seetzeni is active, grows and reproduces in the month following the torrential winter rains. Thereafter, these snails estivate until the following year's rains. By experimental supplementation of water in the field, we examined the ability of these snails to alter their life histories. Specifically, we measured changes in feeding activity, overall activity levels, movement patterns and reproductive output. We found that snails on artificially-watered plots stopped feeding on higher plants and fed on soil-surface algae growing as a result of increased water availability. Most snails moved onto the watered plots, and reduced the total amount of movement once there, in response to increased food availability. Snails on the watered plots grew more and remained active for a longer period into the summer months than did snails on control plots. However, we found little evidence for the ability of these snails to alter their reproductive outputs in response to increased water (and hence food) availability. There was also no carry-over effect of high water availability in one season enhancing reproduction in the following season. We postulate that the low level of phenotypic plasticity recorded here is a consequence of an adaptive conservative strategy to maintain a high reproductive output under extremely unpredictable desert conditions.     

八种杀虫剂对韭菜迟眼蕈蚊发育和繁殖的亚致死效应

用胃毒触杀法确定8种常用杀虫剂对韭菜迟眼蕈蚊Bradysiaodoriphaga 3龄幼虫的LC₂₀和LC₅₀剂量后,以其LC₂₀分别处理3龄幼虫,将存活幼虫正常饲养,测定该剂量药剂对韭菜迟眼蕈蚊化蛹率、蛹重、羽化率,成虫存活率、雌雄比、单雌产卵量及卵孵化率等指标的影响。结果表明:毒死蜱等4种药剂的影响较显著,其影响程度由高到低依次为毒死蜱＞辛硫磷＞阿维菌素溴虫腈;而另4种药剂丙硫克百威、灭多威、丁硫克百威和吡虫啉则影响不显著。此外也测定了用后4种药剂LC₅₀剂量处理的存活试虫的上述生物学指标,结果表明：除化蛹率和羽化率较对照降低且差异显著外,其他生物学指标与对照处理无明显变化。