Predominance of outcrossing in Lymnaea stagnalis despite low apparent fitness costs of self-fertilization

We have quantified the natural mating system in eight populations of the simultaneously hermaphroditic aquatic snail Lymnaea stagnalis, and studied the ecological and genetic forces that may be directing mating system evolution in this species. We investigated whether the natural mating system can be explained by the availability of mates, by the differential survival of self- and cross-fertilized snails in nature, and by the effects of mating system on parental fecundity and early survival. The natural mating system of L. stagnalis was found to be predominantly cross-fertilizing. Density of snails in the populations had no relationship with the mating system, suggesting that outcrossing rates are not limited by mate availability at the population densities observed. Contrary to expectations for outcrossing species, we detected no evidence for inbreeding depression in survival in nature with inferential population genetic methods. Further, experimental manipulations of mating system in the laboratory revealed that self-fertilization had no effect on parental fecundity, and only minor effects on offspring survival. Predominance of cross-fertilization despite low apparent fitness costs of self-fertilization is at odds with the paradigm that high self-fertilization depression is necessary for maintenance of cross-fertilization in self-compatible hermaphrodites.     

BEST OF BOTH WORLDS? ASSOCIATION BETWEEN OUTCROSSING AND PARASITE LOADS IN A SELFING FISH

Mixed-mating strategies (i.e., intermediate levels of self-fertilization and outcrossing in hermaphrodites) are relatively common in plants and animals, but why self-fertilization (selfing) rates vary so much in nature has proved difficult to explain. We tested the hypothesis that parasites help maintain mixed-mating using a partially selfing fish (Kryptolebias marmoratus) as a model. We show that outcrossed progeny in the wild are genetically more diverse and less susceptible to multiple parasite infections than their selfed counterparts. Given that outcrossing in K. marmoratus can only be attained by male-hermaphrodite matings, our data provide an explanation for the coexistence of males and hermaphrodites in androdioecious species where hermaphrodites are unable to outcross among themselves. Moreover, our study provides evidence that parasites contribute to maintaining mixed-mating in a natural animal population.     

Multigenerational inbreeding in <Emphasis Type="Italic">Succisa pratensis</Emphasis>: Effects on fitness components

We examined the effects of repeated inbreeding on fitness components of the long-lived perennial Succisa pratensis (Dipsacaceae). Plants from six populations differing in size were used to establish lines with expected inbreeding coefficients f of 0, 0.5 and 0.75. The effects of different inbreeding levels were measured for seed set, seed mass, percentage germination and seedling relative growth rate. Seed set decreased following one generation of inbreeding and seedling growth rate decreased after two generations of inbreeding. Our study indicated that the mutational load is difficult to purge and that continued inbreeding tends to affect important traits in S. pratensis. Although the partial dominance hypothesis for inbreeding depression seems to account for the results, the overdominance hypothesis cannot be ruled out completely. Overall, we conclude that the response of a long-lived plant, such as S. pratensis, to repeated inbreeding does not differ from that of other plant species with shorter life spans, surely because the mechanisms that account for inbreeding depression are universal for all plant species.     

Somatic deleterious mutation rate in a woody plant: estimation from phenotypic data

We conducted controlled crosses in populations of the long-lived clonal shrub,Vaccinium angustifolium (lowbush blueberry) to estimate inbreeding depressionand mutation parameters associated with somatic deleterious mutation. Inbreedingdepression level was high, with many plants failing to set fruit after self-pollination.We also compared fruit set from autogamous pollinations (pollen collected from within thesame inflorescence) with fruit set from geitonogamous pollinations (pollen collected fromthe same plant but from inflorescences separated by several meters of branch growth). Thedifference between geitonogamous versus autogamous fitness within single plants isreferred to as ‘autogamy depression'' (AD). AD can be caused by somaticdeleterious mutation. AD was significantly different from zero for fruit set. We developeda maximum-likelihood procedure to estimate somatic mutation parameters from AD, andapplied it to geitonogamous and autogamous fruit set data from this experiment. We inferthat, on average, approximately three sublethal, partially dominant somatic mutationsexist within the crowns of the plants studied. We conclude that somatic mutation in thiswoody plant results in an overall genomic deleterious mutation rate that exceeds the ratemeasured to date for annual plants. Some implications of this result for evolutionarybiology and agriculture are discussed.     

HOW ARE DELETERIOUS MUTATIONS PURGED? DRIFT VERSUS NONRANDOM MATING

Accumulation of deleterious mutations has important consequences for the evolution of mating systems and the persistence of small populations. It is well established that consanguineous mating can purge a part of the mutation load and that lethal mutations can also be purged in small populations. However, the efficiency of purging in natural populations, due to either consanguineous mating or to reduced population size, has been questioned. Consequences of consanguineous mating systems and small population size are often equated under \"inbreeding\" because both increase homozygosity, and selection is though to be more efficient against homozygous deleterious alleles. I show that two processes of purging that I call \"purging by drift\" and \"purging by nonrandom mating\" have to be distinguished. Conditions under which the two ways of purging are effective are derived. Nonrandom mating can purge deleterious mutations regardless of their dominance level, whereas only highly recessive mutations can be purged by drift. Both types of purging are limited by population size, and sharp thresholds separate domains where purging is either effective or not. The limitations derived here on the efficiency of purging are compatible with some experimental studies. Implications of these results for conservation and evolution of mating systems are discussed.     

A new theory for the evolution of polyandry as a means of inbreeding avoidance

We propose a novel theory for the evolution of polyandry driven by genetic benefits to females whose offspring interbreed. In species with an ecology characterized by frequent colonization of new habitat patches, consanguineous matings may be common during the early stages of colonization, but genetic diversity may grow as new colonizers arrive. We show that with levels of inbreeding depression similar to those found in predominantly inbreeding populations, a polyandrous female can benefit her descendants since matings among her brood are mainly between half siblings rather than full siblings. We examine the invasion by a polyandrous phenotype using explicit genetic models in which costs of inbreeding are themselves subject to selection. In common with other models of inbreeding, we find that underlying high levels of inbreeding tend to purge deleterious recessive alleles, and hence these are unlikely to maintain sufficient inbreeding depression to favour polyandry. However, if costs of inbreeding are due to overdominance, biologically realistic levels of inbreeding depression result in genetic benefits large enough to favour polyandry provided it is not too costly. The potential significance of polyandry as a mechanism to reduce inbreeding in grandchildren will depend upon the genetic basis of inbreeding depression in natural, inbreeding populations.     

Temporal change in inbreeding depression in life‐history traits in captive populations of guppy (Poecilia reticulata): evidence for purging?

Inbreeding depression, which generally affects the fitness of small populations, may be diminished by purging recessive deleterious alleles when inbreeding persists over several generations. Evidence of purging remains rare, especially because of the difficulties of separating the effects of various factors affecting fitness in small populations. We compared the expression of life-history traits in inbred populations of guppy (Poecilia reticulata) with contemporary control populations over 10 generations in captivity. We estimated inbreeding depression as the difference between the two types of populations at each generation. After 10 generations, the inbreeding coefficient reached a maximum value of 0.56 and 0.16 in the inbred and control populations, respectively. Analysing changes in the life-history traits across generations showed that inbreeding depression in clutch size and offspring survival increased during the first four to six generations in the populations from the inbred treatment and subsequently decreased as expected if purging occurred. Inbreeding depression in two other traits was weaker but showed similar changes across generations. The loss of six populations in the inbred treatment indicates that removal of deleterious alleles also occurred by extinction of populations that presumably harboured high genetic load.