The adaptive accuracy of flowers: measurement and microevolutionary patterns

Background and Aims

From Darwin''s time onward, biologists have thought about adaptation as evolution toward optimal trait values, but they have not usually assessed the relative importance of the distinct causes of deviations from optima. This problem is investigated here by measuring adaptive inaccuracy (phenotypic deviation from the optimum), using flower pollination as an adaptive system.

Methods

Adaptive accuracy is shown to have at least three distinct components, two of which are optimality (deviation of the mean from the optimum) and precision (trait variance). We then describe adaptive accuracy of both individuals and populations. Individual inaccuracy comprises the deviation of the genotypic target (the mean phenotype of a genotype grown in a range of environments) from the optimum and the phenotypic variation around that genotypic target (phenotypic imprecision). Population inaccuracy has three basic components: deviation of the population mean from the optimum, variance in the genotypic targets and phenotypic imprecision. In addition, a fourth component is proposed, namely within-population variation in the optimum. These components are directly estimable, have additive relationships, and allow exploration of the causes of adaptive inaccuracy of both individuals and populations. Adaptive accuracy of a sample of flowers is estimated, relating floral phenotypes controlling pollen deposition on pollinators to adaptive optima defined as the site most likely to get pollen onto stigmas (male inaccuracy). Female inaccuracy is defined as the deviation of the position of stigma contact from the expected location of pollen on pollinators.

Key Results

A surprising amount of variation in estimated accuracy within and among similar species is found. Some of this variation is generated by developmental changes in positions of stigmas or anthers during anthesis (the floral receptive period), which can cause dramatic change in accuracy estimates. There seem to be trends for higher precision and accuracy in flowers with higher levels of integration and dichogamy (temporal separation of sexual functions), and in those that have pollinators that are immobile (or immobilized) during pollen transfer. Large deviations from putative adaptive optima were observed, and these may be related to the effects of conflicting selective pressures on flowers, such as selection against self-pollination promoting herkogamy (spatial separation of pollen and stigmas).

Conclusions

Adaptive accuracy is a useful concept for understanding the adaptive significance of phenotypic means and variances of floral morphology within and among populations and species. Estimating and comparing the various components of adaptive accuracy can be particularly helpful for identifying the causes of inaccuracy, such as conflicting selective pressures, low environmental canalization and developmental instability.Key words: Adaptive accuracy, Collinsia, Dalechampia, fitness, floral precision, Linum, optimality, pollination, Stylidium     

Differences in pollinator faunas may generate geographic differences in floral morphology and integration in Narcissus papyraceus (Amaryllidaceae)

Pollinators may generate selective pressures that affect covariation patterns of multiple traits as well as the mean values of single floral morphological traits. Berg predicted that flowers pollinated by animals whose morphology closely matches the flower's shape will be phenotypically more integrated (tighter correlation of flower traits) than will flowers pollinated by animals not closely fitting the floral morphology. We tested this hypothesis by comparing, in the Strait of Gibraltar region (south Spain, northern Morocco), populations of Narcissus papyraceus that have geographical differences in pollinator faunas. Long-tongued, nectar-feeding moths dominate the pollinator faunas of those populations close to the Strait of Gibraltar, whereas short-tongued, pollen-feeding syrphid flies dominate in peripheral populations farther from the Strait. Populations pollinated by moths and flies differed in the mean values of several floral traits, consistent with the evolution of regional pollination ecotypes. Populations pollinated by moths showed stronger intercorrelation (floral integration) than populations pollinated by hoverflies. Moth-pollinated populations also showed less variation in flower traits than vegetative traits, and this difference was stronger than in fly-pollinated populations. Thus, the pattern of differences in the phenotypic architecture of the Narcissus flowers is consistent with the hypothesis that populations have responded to different selective pressures generated by different pollinators. These data also supported most of the specific predictions of Berg's hypotheses about integration and modularity.     

EXPERIMENTAL ASSESSMENT OF REPRODUCTIVE INTERACTIONS BETWEEN SYMPATRIC ASTER AND ERIGERON (ASTERACEAE) IN INTERIOR ALASKA

Erigeron glabellus and Aster sibiricus have similar flowers, share pollinators, but bloom sequentially in interior Alaska. Both species depend on insect pollination for seed set: the Erigeron is self-incompatible, and the Aster is apparently self-compatible but allogamous. To test the hypothesis that sequential blooming is maintained by natural selection generated by reproductive interference, we manipulated the flowering time of Erigeron, forcing it to bloom simultaneously with Aster, and measured female fecundity in both species. We found no evidence of reduced female fecundity in either species caused by the presence of the sympatric “competitor” or by artificial pollination with the heterospecific pollen prior to conspecific pollination. Two-species mixtures of simultaneously blooming Aster and Erigeron experienced significant interspecific visitation, which may, under natural conditions, cause loss of pollen to alien stigmas and depressed male fecundity, at least in Erigeron. We found no evidence that sequential blooming in Erigeron and Aster is maintained by depressed female fecundity through pollinator sharing. If sequential blooming is maintained by natural selection, it seems more likely to be the result of selection generated by depressed male fitness through pollen loss to alien stigmas.     

Downregulation of calcineurin activity in cervical carcinoma

Background  

Calcineurin (CaN) is an important serine-threonine phosphatase (PP2B), which plays a crucial role in calcium-calmodulin mediated signal transduction events. Calcineurin has been implicated in pathogenesis of various diseases cardiac hypertrophy, diabetic neuropathy and Alzheimer's, however its role in neoplasia remains unclear.     

Basketball Teams as Strategic Networks

We asked how team dynamics can be captured in relation to function by considering games in the first round of the NBA 2010 play-offs as networks. Defining players as nodes and ball movements as links, we analyzed the network properties of degree centrality, clustering, entropy and flow centrality across teams and positions, to characterize the game from a network perspective and to determine whether we can assess differences in team offensive strategy by their network properties. The compiled network structure across teams reflected a fundamental attribute of basketball strategy. They primarily showed a centralized ball distribution pattern with the point guard in a leadership role. However, individual play-off teams showed variation in their relative involvement of other players/positions in ball distribution, reflected quantitatively by differences in clustering and degree centrality. We also characterized two potential alternate offensive strategies by associated variation in network structure: (1) whether teams consistently moved the ball towards their shooting specialists, measured as “uphill/downhill” flux, and (2) whether they distributed the ball in a way that reduced predictability, measured as team entropy. These network metrics quantified different aspects of team strategy, with no single metric wholly predictive of success. However, in the context of the 2010 play-offs, the values of clustering (connectedness across players) and network entropy (unpredictability of ball movement) had the most consistent association with team advancement. Our analyses demonstrate the utility of network approaches in quantifying team strategy and show that testable hypotheses can be evaluated using this approach. These analyses also highlight the richness of basketball networks as a dataset for exploring the relationships between network structure and dynamics with team organization and effectiveness.     

Does multiple paternity affect seed mass in angiosperms? An experimental test in Dalechampia scandens

Flowers fertilized by multiple fathers may be expected to produce heavier seeds than those fertilized by a single father. However, the adaptive mechanisms leading to such differences remain unclear, and the evidence inconsistent. Here, we first review the different hypotheses predicting an increase in seed mass when multiple paternity occurs. We show that distinguishing between these hypotheses requires information about average seed mass, but also about within‐fruit variance in seed mass, bias in siring success among pollen donors, and whether siring success and seed mass are correlated. We then report the results of an experiment on Dalechampia scandens (Euphorbiaceae), assessing these critical variables in conjunction with a comparison of seed mass resulting from crosses with single vs. multiple pollen donors. Siring success differed among males when competing for fertilization, but average seed mass was not affected by the number of fathers. Furthermore, paternal identity explained only 3.8% of the variance in seed mass, and siring success was not correlated with the mass of the seeds produced. Finally, within‐infructescence variance in seed mass was not affected by the number of fathers. These results suggest that neither differential allocation nor sibling rivalry has any effect on the average mass of seeds in multiply sired fruits in D. scandens. Overall, the limited paternal effects observed in most studies and the possibility of diversification bet hedging among flowers (but not within flowers), suggest that multiple paternity within fruits or infructescence is unlikely to affect seed mass in a large number of angiosperm species.