Kenneth Wilson

Noise and determinism in synchronized sheep dynamics

A major debate in ecology concerns the relative importance of intrinsic factors and extrinsic environmental variations in determining population size fluctuations. Spatial correlation of fluctuations in different populations caused by synchronous environmental shocks,, is a powerful tool for quantifying the impact of environmental variations on population dynamics,. However, interpretation of synchrony is often complicated by migration between populations,. Here we address this issue by using time series from sheep populations on two islands in the St Kilda archipelago. Fluctuations in the sizes of the two populations are remarkably synchronized over a 40-year period. A nonlinear time-series model shows that a high and frequent degree of environmental correlation is required to achieve this level of synchrony. The model indicates that if there were less environmental correlation, population dynamics would be much less synchronous than is observed. This is because of a threshold effect that is dependent on population size; the threshold magnifies random differences between populations. A refined model showsthat part of the required environmental synchronicity can be accounted for by large-scale weather variations. These results underline the importance of understanding the interaction between intrinsic and extrinsic influences on population dynamics.

Dominant rams lose out by sperm depletion

A waning success in siring counters a rams high score in competition for ewes.

Flexible diet choice offsets protein costs of pathogen resistance in a caterpillar

Mounting effective resistance against pathogens is costly in terms of energy and nutrients. However, it remains unexplored whether hosts can offset such costs by adjusting their dietary intake so as to recoup the specific resources involved. We test this possibility by experimentally challenging caterpillars (Spodoptera littoralis) with a highly virulent entomopathogen (nucleopolyhedrovirus), under dietary regimes varying in the content of protein and digestible carbohydrate. We found that dietary protein influenced both resistance to pathogen attack and constitutive immune function to a greater extent than did dietary carbohydrate, indicating higher protein costs of resistance than energy costs. Moreover, when allowed to self-compose their diet, insects surviving viral challenge increased their relative intake of protein compared with controls and those larvae dying of infection, thus demonstrating compensation for protein costs associated with resistance. These results suggest that the change in the hosts nutritional demands to fight infection induces a compensatory shift in feeding behaviour.

Coping with crowds: Density-dependent disease resistance in desert locusts

Parasite transmission generally exhibits some form of positive density dependence. Thus, as population density increases, so too does the per capita risk of becoming infected. Under such circumstances, natural selection should favor individuals that use cues associated with population density to determine the optimal allocation of resources to disease resistance mechanisms. As a consequence, individuals experiencing crowded conditions are predicted to be more resistant to parasites and pathogens than those experiencing low-density conditions. This phenomenon (termed “density-dependent prophylaxis”) [Wilson, K. & Reeson, A. F. (1998) Ecol. Entomol. 23, 100–101] is predicted to be particularly prevalent in outbreak pest species and in species exhibiting density-dependent phase polyphenism, such as the desert locust, Schistocerca gregaria. Here we show that, as predicted, desert locusts reared under crowded conditions are significantly more resistant than solitary locusts to the entomopathogenic fungus, Metarhizium anisopliae var. acridum, a key natural disease of acridids and an important agent in locust and grasshopper biocontrol. Moreover, enhanced pathogen resistance in crowded locusts is associated with elevated antimicrobial activity, but not with any difference in thermal preferences or behavioral fever response. These results have implications for understanding the development and biocontrol of locust plagues.

Costs of resistance: genetic correlations and potential trade‐offs in an insect immune System

Theory predicts that natural selection will erode additive genetic variation in fitness‐related traits. However, numerous studies have found considerable heritable variation in traits related to immune function, which should be closely linked to fitness. This could be due to trade‐offs maintaining variation in these traits. We used the Egyptian cotton leafworm, Spodoptera littoralis, as a model system to examine the quantitative genetics of insect immune function. We estimated the heritabilities of several different measures of innate immunity and the genetic correlations between these immune traits and a number of life history traits. Our results provide the first evidence for a potential genetic trade‐off within the insect immune system, with antibacterial activity (lysozyme‐like) exhibiting a significant negative genetic correlation with haemocyte density, which itself is positively genetically correlated with both haemolymph phenoloxidase activity and cuticular melanization. We speculate on a potential trade‐off between defence against parasites and predators, mediated by larval colour, and its role in maintaining genetic variation in traits under natural selection.

Generalized linear modelling for parasitologists

Typically, the distribution of macroparasites over their host population is highly aggregated and empirically best described by the negative binomial distribution. For parasitologists, this poses a statistical provlem, which is often tackled by log-transforming the parasite data prior to analysis by parametric tests. Here, Ken Wilson and Bryan Grenfell show that this method is particularly prone to type I errors, and highlight a much more powerful and flexible alternative: generalized linear modelling.

Baculovirus resistance in the noctuid Spodoptera exempta is phenotypically plastic and responds to population density

Parasite resistance mechanisms can be costly to maintain. We would therefore predict that organisms should invest in resistance only when it is likely to be required. Insects that show density–dependent phase polyphenism, developing different phenotypes at high and low population densities, have the opportunity to match their levels of investment in resistance with the likelihood of exposure to pathogens. As high population densities often precipitate disease epidemics, the high–density form should be selected to invest relatively more in resistance. We tested this prediction in larvae of the noctuid Spodoptera exempta. Larvae reared at a high density were found to be considerably more resistant to a nuclear polyhedrosis virus than those reared in isolation. A conspicuous feature of the high–density phase of S. exempta and other phase–polyphenic Lepidoptera is cuticular melanization. As melanization is controlled by the phenoloxidase enzyme system, which is also involved in the immune response, this suggests a possible mechanism for increased resistance at high population densities. We demonstrated that melanized S. exempta larvae were more resistant than non–melanized forms, independent of rearing density. We also found that haemolymph phenoloxidase activity was correlated with cuticular melanization, providing further evidence for a link between melanization and immunity. These results suggest that pathogen resistance in S. exempta is phenotypically plastic, and that the melanized cuticles characteristic of the high–density form may be indicative of a more active immune system.

STABILITY AND INSTABILITY IN UNGULATE POPULATIONS: AN EMPIRICAL ANALYSIS

While many populations of large mammals are stable from year to year, some show persistent oscillations associated with high mortality. This article investigates the causes of variation in population stability in ungulates by comparing the contrasting dynamics of two naturally regulated island populations: the Soay sheep population of Hirta (St. Kilda), where numbers fluctuate by 60%-70% every 3 or 4 yr as a result of overcompensatory winter mortality, and the red deer population of the North Block of Rum, where numbers have been stable for over a decade. We suggest that the contrasting dynamics of these two populations are caused by differences in fecundity and maturation rate. Red deer show relatively low fecundity and slow maturation, and increases in population density depress both fecundity and juvenile survival. In contrast, the relatively rapid development of Soay sheep allows them to evade densitydependent effects on reproduction and survival until the population exceeds winter carrying capacity by a substantial margin, subsequently triggering overcompensatory mortality in late winter in all age classes.

A microsatellite polymorphism in the gamma interferon gene is associated with resistance to gastrointestinal nematodes in a naturally-parasitized population of Soay sheep

Free-living Soay sheep (Ovis aries) on the island of Hirta, St Kilda, Scotland, are naturally parasitized by gastrointestinal nematodes, predominantly Teladorsagia circumcincta. In this paper we show that reduced faecal egg counts (FEC) are associated with an allele at a microsatellite locus located in the first intron of the interferon gamma gene (o(IFN)-gamma) in Soay sheep lambs and yearlings, measured at approximately 4 and 16 months of age, respectively. The same allele was also associated with increased T. circumcincta-specific antibody (IgA) in lambs, but not associated significantly in yearlings. Flanking control markers failed to show a significant association with either FEC or IgA. These results suggest that a polymorphic gene conferring increased resistance to gastrointestinal nematode parasites is located at or near the interferon gamma gene, and support previous reports which have mapped a quantitative trait locus (QTL) for resistance to this region in domestic sheep. Our data are consistent with the idea that a functional polymorphism leading to reduced expression or efficacy of (IFN)-gamma could enhance the immune response to gastrointestinal nematodes by favouring the activity of the Th2 cell subset and antibody associated immune mechanisms.

Immunocompetence: a neglected life history trait or conspicuous red herring?

We thank everyone whom directly or indirectly contributed to this review, especially Victor Apanius, Matthew Evans, Bryan Grenfell, Frances Gulland, Bob Lochmiller, Anders Moller, Andrew Read, Ben Sheldon, Mike Siva-Jothy and Simon Verhulst. I.P.F.O. also thanks Paul Harvey for hospitality during his sabbatical and K.W. acknowledges the support of the NERC.