COMPONENTS OF PHENOTYPIC SELECTION: POLLEN EXPORT AND FLOWER COROLLA WIDTH IN IPOMOPSIS AGGREGATA

In the hummingbird-pollinated herb Ipomopsis aggregata, selection through male function during pollination favors wide corolla tubes. We explored the mechanisms behind this selection, using phenotypic selection analysis to compare effects of corolla width on two components of male pollination success, pollinator visit rate and pollen exported per visit. During single visits by captive hummingbirds, flowers with wider corollas exported more pollen, and more dye used as a pollen analogue, to stigmas of recipient flowers. Corolla width was less strongly related to visit rate in the field, and had no direct effect on visit rate after nectar production and corolla length were controlled for. Moreover, the phenotypic selection differential was 80% higher for the effect on pollen exported per visit, suggesting that this is the more important mechanism of selection.     

Direct and indirect responses to selection on pollen size in Brassica rapa L.

Pollen size varies little within angiosperm species, but differs extensively between species, suggesting the action of strong selection. Nevertheless, the potential for genetic responses of pollen size to selection, as determined by additive genetic variance and genetic correlations with other floral traits, has received little attention. To assess this potential, we subjected Brassica rapa to artificial selection for large and small pollen during three generations. This selection caused significant divergence in pollen diameter, with additive genetic effects accounting for over 30% of the observed phenotypic variation in pollen size. Such heritable genetic variation suggests that natural selection could effect evolutionary change in this trait. Selection on pollen size also elicited correlated responses in pollen number (–), flower size (+), style length (+), and ovule number (+), suggesting that pollen size cannot evolve independently. The correlated responses of pollen number, flower size and ovule number probably reflect the genetically determined and physically constrained pattern of resource allocation in B. rapa. In contrast, the positive correlation between pollen size and style length may represent a widespread gametic‐phase disequilibrium in angiosperms that arises from nonrandom fertilization success of large pollen in pistils with long styles.     

Floral nectar production: what cost to a plant?

Floral nectar production is central to plant pollination, and hence to human wellbeing. As floral nectar is essentially a solution in water of various sugars, it is likely a valuable plant resource, especially in terms of energy, with plants experiencing costs/trade-offs associated with its production or absorption and adopting mechanisms to regulate nectar in flowers. Possible costs of nectar production may also influence the evolution of nectar volume, concentration and composition, of pollination syndromes involving floral nectar, and the production of some crops. There has been frequent agreement that costs of floral nectar production are significant, but relevant evidence is scant and difficult to interpret. Convincing direct evidence comes from experimental studies that relate either enhanced nectar sugar production (through repeated nectar removal) to reduced ability to produce seeds, or increased sugar availability (through absorption of additional artificial nectar) to increased seed production. Proportions of available photosynthate allocated by plants to nectar production may also indicate nectar cost. However, such studies are rare, some do not include treatments of all (or almost all) flowers per plant, and all lack quantitative cost–benefit comparisons for nectar production. Additional circumstantial evidence of nectar cost is difficult to interpret and largely equivocal. Future research should repeat direct experimental approaches that relate reduced or enhanced nectar sugar availability for a plant with consequent ability to produce seeds. To avoid confounding effects of inter-flower resource transfer, each plant should experience a single treatment, with treatment of all or almost all flowers per plant. Resource allocation by plants, pathways used for resource transfer, and the locations of resource sources and sinks should also be investigated. Future research should also consider extension of nectar cost into other areas of biology. For example, evolutionary models of nectar production are rare but should be possible if plant fitness gains and costs associated with nectar production are expressed in the same currency, such as energy. It should then be possible to understand observed nectar production for different plant species and pollination syndromes involving floral nectar. In addition, potential economic benefits should be possible to assess if relationships between nectar production and crop value are evaluated.     

Divergent selection on,but no genetic conflict over,female and male timing and rate of reproduction in a human population

The sexes often have different phenotypic optima for important life-history traits, and because of a largely shared genome this can lead to a conflict over trait expression. In mammals, the obligate costs of reproduction are higher for females, making reproductive timing and rate especially liable to conflict between the sexes. While studies from wild vertebrates support such sexual conflict, it remains unexplored in humans. We used a pedigreed human population from preindustrial Finland to estimate sexual conflict over age at first and last reproduction, reproductive lifespan and reproductive rate. We found that the phenotypic selection gradients differed between the sexes. We next established significant heritabilities in both sexes for all traits. All traits, except reproductive rate, showed strongly positive intersexual genetic correlations and were strongly genetically correlated with fitness in both sexes. Moreover, the genetic correlations with fitness were almost identical in men and women. For reproductive rate, the intersexual correlation and the correlation with fitness were weaker but again similar between the sexes. Thus, in this population, an apparent sexual conflict at the phenotypic level did not reflect an underlying genetic conflict over the studied reproductive traits. These findings emphasize the need for incorporating genetic perspectives into studies of human life-history evolution.