Evolvability and constraint in the evolution of three-dimensional flower morphology.

PREMISE: Flower phenotypes evolve to attract pollinators and to ensure efficient pollen transfer to and from the bodies of pollinators or, in self-compatible bisexual flowers, between anthers and stigmas. If functionally interacting traits are genetically correlated, response to selection may be subject to genetic constraints. Genetic constraints can be assessed by quantifying standing genetic variation in (multivariate) phenotypic traits and by asking how much the available variation is reduced under specific assumptions about phenotypic selection on functionally interacting and genetically correlated traits. METHODS: We evaluated multivariate evolvability and potential genetic constraints underlying the evolution of the three-dimensional structure of Dalechampia blossoms. First, we used data from a greenhouse crossing design to estimate the G matrix for traits representing the relative positions of male and female sexual organs (anthers and stigmas) and used the G matrix to ask how genetic variation is distributed in multivariate space. To assess the evolutionary importance of genetic constraints, we related standing genetic variation across phenotypic space to evolutionary divergence of population and species in the same phenotypic directions. RESULTS: Evolvabilities varied substantially across phenotype space, suggesting that certain traits or trait combinations may be subject to strong genetic constraint. Traits involved functionally in flower-pollinator fit and autonomous selfing exhibited considerable independent evolutionary potential, but population and species divergence tended to occur in phenotypic directions associated with greater-than-average evolvability. CONCLUSIONS: These results are consistent with the hypothesis that genetic constraints can hamper joint trait evolution towards optimum flower-pollinator fit and optimum self-pollination rates.

Fitness costs of delayed pollination in a mixed-mating plant.

BACKGROUND AND AIMS: To predict the evolutionary consequences of pollinator declines, we need to understand the evolution of delayed autonomous self-pollination, which is expected to evolve as a mechanism of reproductive assurance when cross-pollination becomes unreliable. This involves estimating the costs of increased levels of selfing as well as those associated with floral senescence. METHODS: We studied the mechanisms and costs of delayed self-pollination in the mixed-mating vine Dalechampia scandens (Euphorbiaceae) by first assessing among-population variation in herkogamy and dichogamy, which together determine the rate and timing of autonomous self-pollination. We then tested whether floral longevity responds plastically to delayed pollination. Finally, we assessed the costs of delayed self-pollination in terms of seed number and size, explicitly separating inbreeding depression from effects of floral senescence. KEY RESULTS: Herkogamy varied extensively, while variation in dichogamy was more limited. Unpollinated blossoms increased their longevity, but seed quantity and quality decreased with increasing delays in pollination, independently of inbreeding depression. CONCLUSIONS: In D. scandens, earlier autonomous selfing is facilitated by reduced herkogamy rather than reduced protogyny, providing reproductive assurance while maintaining the possibility for outcrossing events. Effective early autonomous self-pollination may evolve under reduced cross-pollination reliability in response to costs associated with floral senescence.

Quantitative and qualitative consequences of reduced pollen loads in a mixed-mating plant.

Greater pollination intensity can enhance maternal plant fitness by increasing seed set and seed quality as a result of more intense pollen competition or enhanced genetic sampling. We tested experimentally these effects by varying the pollen load from a single pollen donor on stigmas of female flowers of Dalechampia scandens (Euphorbiaceae) and measuring the effects on seed number and seed mass. Seed set increased rapidly with pollen number at low to moderate pollen loads, and a maximum set of three seeds occurred with a mean pollen load of 19 pollen grains. We did not detect a trade-off between the number of seeds and seed mass within a fruit. Seed mass increased with increasing pollen load, supporting the hypothesis of enhanced seed quality via increased pollen-competition intensity or genetic sampling. These results suggest that maternal fitness increases with larger pollen loads, even when the fertilization success is already high. Our results further highlight the importance of high rates of pollen arrival onto stigmas, as mediated by reliable pollinators. Comparing the pollen-to-seed response curve obtained in this experiment with those observed in natural populations suggests that pollen limitation may be more severe in natural populations than predicted from greenhouse studies. These results also indicate that declines in pollinator abundance may decrease plant fitness through lowered seed quality before an effect on seed set is detected.