Despite the importance of adaptation in shaping biological diversity over many generations, little is known about populations’ capacities to adapt at any particular time. Theory predicts that a population's rate of ongoing adaptation is the ratio of its additive genetic variance for fitness, VAfalse(Wfalse), to its mean absolute fitness, W¯. We conducted a transplant study to quantify W¯ and standing VAfalse(Wfalse) for a population of the annual legume Chamaecrista fasciculata in one field site from which we initially sampled it and another site where it does not currently occur naturally. We also examined genotype‐by‐environment interactions, G × E, as well as its components, differences between sites in VAfalse(Wfalse) and in rank of breeding values for fitness. The mean fitness indicated population persistence in both sites, and there was substantial VAfalse(Wfalse) for ongoing adaptation at both sites. Statistically significant G × E indicated that the adaptive process would differ between sites. We found a positive correlation between fitness of genotypes in the “home” and “away” environments, and G × E was more pronounced as the life‐cycle proceeds. This study exemplifies an approach to assessing whether there is sufficient VAfalse(Wfalse) to support evolutionary rescue in populations that are declining.
Plant-pollinator interactions promote the evolution of floral traits that attract pollinators and facilitate efficient pollen transfer. The spatial separation of sex organs, herkogamy, is believed to limit sexual interference in hermaphrodite flowers. Reverse herkogamy (stigma recessed below anthers) and long, narrow corolla tubes are expected to promote efficiency in male function under hawkmoth pollination. We tested this prediction by measuring selection in six experimental arrays of Polemonium brandegeei, a species that displays continuous variation in herkogamy, resulting in a range of recessed to exserted stigmas. Under glasshouse conditions, we measured pollen removal and deposition, and estimated selection gradients (β) through female fitness (seeds set) and male fitness (siring success based on six polymorphic microsatellite loci). Siring success was higher in plants with more nectar sugar and narrow corolla tubes. However, selection through female function for reverse herkogamy was considerably stronger than was selection through male function. Hawkmoths were initially attracted to larger flowers, but overall preferred plants with reverse herkogamy. Greater pollen deposition and seed set also occurred in reverse herkogamous plants. Thus, reverse herkogamy may be maintained by hawkmoths through female rather than male function. Further, our results suggest that pollinator attraction may play a considerable role in enhancing female function.
Production of multiple flowers in inflorescences allows the reproductive phenotypes of individual plants to include systematic among-flower variation, which could be adaptive. Systematic trait variation within inflorescences could arise from resource competition among flowers, or be a developmentally determined feature of flower position, regardless of resource dynamics. The latter, architectural effect typically manifests as continuous floral variation within inflorescences. For architectural effects to be adaptive, floral trait variation among individuals must covary with reproductive performance and be heritable. However, heritability and phenotypic selection on gradients of variation cannot be estimated readily with traditional statistical approaches. Instead, we advocate and illustrate the application of two functional data analysis techniques with observations of Delphinium glaucum (Ranunculaceae). To demonstrate the parameters-as-data approach we quantify heritability of variation in anthesis rate, as represented by the regression coefficient relating daily anthesis rate to inflorescence age. SNP-based estimates detected significant heritability (h 2 ¼ 0.245) for declining anthesis rate within inflorescences. Functional regression was used to assess phenotypic selection on anthesis rate and a floral trait (lower sepal length). The approach used spline curves that characterize within-inflorescence variation as functional predictors of a plant's fruit set. Selection on anthesis rate varied with inflorescence age and the duration of an individual's anthesis period. Lower sepal length experienced positive selection for basal and distal flowers, but negative selection for central flowers. These results illustrate the utility and power of functional-data analyses for studying architectural effects and specifically demonstrate that these effects are subject to natural selection and hence adaptive.
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