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One of the leading hypotheses for the maintenance of sexual reproduction is the Red Queen hypothesis. The underlying premise of the Red Queen hypothesis is that parasites rapidly evolve to infect common host genotypes. This response by parasites could result in the long-term maintenance of genetic variation and may favor sexual reproduction over asexual reproduction. The underlying ideas present a wonderful microcosm for teaching evolution. Here I present the reasons for why sex is anomalous for evolutionary theory, the rationale underlying the Red Queen hypothesis, and some empirical studies of the Red Queen hypothesis using a freshwater snail. The empirical results are consistent with the Red Queen hypothesis. In addition, the distribution of sexual and asexual reproduction in the snail leads naturally to thinking about coevolution in a geographic mosaic of parasite-mediated natural selection.

Host-parasite interactions that result in host castration are evolutionarily similar to predator-prey interactions because both interactions terminate reproduction for the host or prey. Yet, host-parasite interactions differ from predator-prey interactions in that infected hosts remain alive and potentially can make adjustments to their life-history strategy before castration is complete. Here we exposed juvenile snails (Potamopyrgus antipodarum) to infection by a digenetic trematode (Microphallus sp.) in order to determine whether: (1) pre-reproductive individuals could be infected, (2) individuals that were exposed to infection shifted resources to early reproduction (fecundity compensation), and (3) infected individuals exhibit altered growth rates relative to uninfected individuals. We found that juveniles are susceptible to infection; hence P. antipodarum could be selected for earlier maturation in populations where the risk of infection is high. We also found that fecundity compensation does not occur in this snail. Finally, we found that Microphallus-infected snails exhibit altered growth rates; individuals infected as juveniles have lower growth rates and are smaller than uninfected snails. These results suggest that growth is altered by infection of a trematode parasite but reproduction in uninfected snails is not induced by exposure to trematode eggs.

Software reuse is widely believed to be a key to improving software productivity and quality in conventional software. In expert systems, much of the knowledge has been compiled (i.e., compressed and restricted into effective procedures) and this makes reusability difficult. One of the issues in modeling expert systems for enhanced reusability is capturing explicity the underlying problem solving designs. Principled knowledge representation schemes have been used to model components of complex software systems. However, the potential for applying these principled modeling techniques for explicitly capturing the problem solving designs of expert systems has not been fully explored. To overcome this omission, we use an Artificial Intelligence knowledge representation scheme for developing an ontology of the software components to facilitate their classification and retrieval. The application of our ontological approach is of both theoretical and practical significance. This method facilitates the reuse of high-level design. We illustrate the application of principled domain modeling using two real world applications of knowledge-based systems.

MANY species exhibit colour polymorphisms associated with alternative male reproductive strategies, including territorial males and 'sneaker males' that behave and look like females1–3. The prevalence of multiple morphs is a challenge to evolutionary theory because a single strategy should prevail unless morphs have exactly equal fitness4,5 or a fitness advantage when rare6,7. We report here the application of an evolutionary stable strategy model to a three-morph mating system in the side-blotched lizard. Using parameter estimates from field data, the model predicted oscillations in morph frequency, and the frequencies of the three male morphs were found to oscillate over a six-year period in the field. The fitnesses of each morph relative to other morphs were non-transitive in that each morph could invade another morph when rare, but was itself invadable by another morph when common. Concordance between frequency-dependent selection and the among-year changes in morph fitnesses suggest that male interactions drive a dynamic 'rock–paper–scissors' game7.

We review the spatial and temporal correlates of uniparental reproduction in freshwater snails as they pertain to the ecological hypotheses for the maintenance of biparental sex. The biogeographic evidence from two species (Potamopyrgus antipodarum andBulinus truncatus) presently supports the Red Queen hypothesis that biparental reproduction is selected as a way to reduce the risk to progeny of parasite attack. Uniparental reproduction in these species is associated with low levels of infection by parasites (castrating digenetic trematodes), suggesting that parthenogenesis or self-fertilization can replace cross-fertilization when the risk of infection is low. In addition, inB. truncatus, the opportunity for cross-fertilization coincides with the season in which parasite attack is highest. In a third species (Campeloma decisum), parthenogenetic reproduction is correlated with latitude and the presence of a non-castrating trematode that may prevent cross-fertilization; these patterns suggest that parthenogenesis has been selected as a mechanism to assure reproduction. Finally, we discuss the spotty taxonomic distribution of parthenogenetic species.

Larval digenetic trematodes commonly castrate their first intermediate hosts, and should therefore impose strong selection on the timing and mode of host reproduction. Here we examine spatial variation in infection by trematodes in the freshwater snail Potamopyrgus antipodarum. Snails were collected at 11 different sites at Lake Alexandrina on the South Island of New Zealand from transects that ran perpendicular to the shore and across several different habitat types (from 0 to 8 m deep). Logistic regression was used to analyze the relationships between the frequency of trematode infection and snail size, habitat type, and transect location. On average, the probability of infection increased 3.3 times with each 1 mm increase in shell length. Prevalence of infection by the most common species of trematode, Microphallus sp., was highest in the shallow-water habitats where its final hosts (waterflow) spend most of their time. Prevalence of infection by another parasite, Telogaster ophistorchis (final host: eels) increased with depth, but because Microphallus was much more common, total infection by all trematodes decreased with depth. The effects of transect location were minor for Telogaster, but there was significant variation in Microphallus prevalence among transects, especially in the shore-bank habitat. Taken together, these results suggest that the risk of infection is spatially variable, but generally higher in shallow-water habitats, which may explain the greater frequency of sexual individuals as well as earlier reproduction among individuals near shore.

The quadrupole moment for the πg 9/2⊗νh 11/2 band in the 130>Pr nucleus has been measured using the Doppler-shift attenuation method. A centroid shift analysis was carried out and a value of Q0=6.1±0.4 eb, corresponding to an axial prolate deformation of β2=0.35(3), has been determined. This is the first direct experimental confirmation of the deformation-driving character of the πg 9/2 orbital in an odd-odd nucleus in the A≃135 superdeformed region.

THE Red Queen hypothesis for the maintenance of biparental sexual reproduction suggests that, for species locked in revolutionary struggles with biological enemies, the production of variable progeny compensates for the genetic or ecological disadvantages of sex1–7. The advantage of sex and recombination under this hypothesis stems from the production of rare phenotypes, which are expected to be more likely to escape infection or predation by coevolved biological enemies. Like many evolutionary hypotheses, the Red Queen hypothesis is difficult to test directly, but its assumptions and predictions can be evaluated7–18. The most critical assumption is that biological enemies will disproportionately attack the most common phenotype19, 20. In this study of parasite loads of coexisting sexual and clonal fish, we find empirical support for this assumption.

Background Female mate choice may be adaptive when males exhibit heritable genetic variation at loci encoding resistance to infectious disease. The Hamilton-Zuk hypothesis predicts that females should assess the genetic quality of males by monitoring traits that indicate health and vigor (condition-dependent choice, or CD). Alternatively, some females may employ a more direct method of screening and select mates based on the dissimilarity of alleles at the major histocompatibility loci (we refer to this as opposites-attract, or OA). Empirical studies suggest that both forms of mate choice exist, but little is known about the potential for natural selection to shape the two strategies in nature.

Nature 367, 554-557 (1994) DURING the production process, the contrast in shading in Figs 1 and 2 of this letter was lost; in addition, parts a and b of Fig. 3 were transposed. These figures should have appeared as they are shown here. The first sentence of each legend is given for guidance. FIG. 1 Results from computer simulations in which sexual populations were challenged by asexual lineages in the presence of coevolving parasites.

Field experiments were conducted in order to determine the potential for desiccation and predation to mediate the effect of mussels (Brachidontes semilaevis) on barnacles (Chthamalus anisopoma) in the highly seasonal northern Gulf of California. We did this by removing both mussels and a common mussel predator (Morula ferruginosa: Gastropoda) and by spraying selected sites with sea water during summertime spring low tides. We also determined the effect of crowding on resistance to desiccation in barnacles, and the effect of barnacles on colonization by mussels. The mussel-barnacle community was not affected by keeping experimental quadrats damp during daytime low tides throughout the summer. Exposure to summertime low tides, however, did affect the survivorship of isolated, but not crowded, barnacles; and barnacle clumps enhanced the recruitment of mussels. Hence crowding in barnacles had a positive effect on both barnacle survivorship and mussel recruitment. Morula had a negative effect on mussel density, and mussels had a negative effect on barnacle density. The effect of Morula on barnacle density was positive, presumably due to its selective removal of mussels. These results suggest an indirect mutualism between barnacles and the gastropod predator, because barnacles attract settlement or enhance the survival of mussels, and the predator reduces the competitive effect of mussels on barnacles.

Phenotypic models of selection are used to determine the effect of facultative parthenogenesis on the production of males in a spatially variable environment when (i) sex determination is under strict genetic control, and (ii) when sex may be environmentally determined. The results show that when sex is under strict genetic control and there is some chance of maturing in isolation, selection favors a female-biased sex ratio. When sex can be environmentally induced by cues which indicate high density, selection favors a mixture of genetic and environmental control, such that half the individuals always become female and the other half become females when isolated and become males when not isolated.

The widespread occurrence of sexual reproduction is an important problem in evolutionary theory, because of the cost of producing males in dioecious organisms and the allocation of limited resources in cosexes1–4. Here I compare the predictions of the major ecological hypotheses on the maintenance of sex by examining male frequency in Potamopyrgus antipodarum, a dioecious freshwater snail with both sexual and parthenogenetic populations5. The results do not-support the view that sex is maintained by a variable physical environment6–9, but they are consistent with the idea that sex is favoured by selection resulting from host–parasite interactions10–14.