Artificial selection shifts flowering phenology and other correlated traits in an autotetraploid herb.

There is mounting evidence that plants are responding to anthropogenic climate change with shifts in flowering phenologies. We conducted a three-generation artificial selection experiment on flowering time in Campanulastrum americanum, an autotetraploid herb, to determine the potential for adaptive evolution of this trait as well as possible costs associated with enhanced or delayed flowering. Divergent selection for earlier and later flowering resulted in a 25-day difference in flowering time. Experiment-wide heritability was 0.31 and 0.23 for the initiation of flowering in early and late lines, respectively. Selection for earlier flowering resulted in significant correlated responses in other traits including smaller size, fewer branches, smaller floral displays, longer fruit maturation times, fewer seeds per fruit and slower seed germination. Results suggest that although flowering time shows the potential to adapt to a changing climate, phenological shifts may be associated with reduced plant fitness possibly hindering evolutionary change.

Population differentiation and hybrid success in Campanula americana: geography and genome size.

Populations within a species may diverge through genetic drift and natural selection. Few studies report on population differentiation in autopolyploids where multiple gene copies and the ratio of cytoplasmic to nuclear genes differ from diploids and may influence divergence. In autotetraploid Campanula americana we created hybrids between populations that differed in geographic proximity and genome size. Differences in genome size (up to 6.5%) did not influence hybrid performance. In contrast, hybrid performance was strongly influenced by population proximity. F1 hybrids between distant populations performed poorly relative to their parents while hybrids between proximate populations outperformed their parents. Outbreeding depression was strongest for juvenile traits. The expression of outbreeding depression often differed between reciprocal hybrids indicating interactions between nuclear and cytoplasmic genes contribute to population differentiation. Because plants were grown under greenhouse conditions, the outbreeding depression was likely due to genetic (underdominance or loss of additive-by-additive epistasis) rather than ecological factors.

Outcrossing rate and inbreeding depression in the herbaceous autotetraploid, Campanula americana.

Polyploidy in angiosperms is frequently associated with an increase in self-compatibility. Self-fertilization can enhance polyploid establishment, and theory predicts reduced inbreeding depression in polyploids relative to diploids. Therefore, we may expect mating systems that promote self-fertilization or mixed-mating in polyploid species. However, few studies have measured polyploid mating systems and inbreeding depression. We report the outcrossing rate and inbreeding depression for Campanula americana, a self-compatible protandrous herb. Allozyme genotypes suggest that C. americana is an autotetraploid with tetrasomic inheritance. We found that the multilocus outcrossing rate, t(m)=0.938, did not differ from unity. This result was unexpected since previous work demonstrated that pollinators frequently move from male- to female-phase flowers on the same plant, that is, geitonogamy. Self and outcross pollinations were conducted for three populations. Offspring were germinated in controlled conditions and grown to maturity in pots in nature. Inbreeding depression was not significant for most seed and germination characters. However, all later life traits except flowering date differed between inbred and outcrossed individuals resulting in a 26% reduction in cumulative fitness for inbred plants. Limited early- and moderate later-life inbreeding depression suggest that it is buffered by the higher levels of heterozygosity found in an autotetraploid. C. americana appears to have a flexible mating system where within flower protandry and/or cryptic self-incompatibility result in a high outcrossing rate when pollinators are abundant, but self-compatibility and limited inbreeding depression maintain reproductive success when mates are limited.

Constraint to adaptive evolution in response to global warming.

We characterized the genetic architecture of three populations of a native North American prairie plant in field conditions that simulate the warmer and more arid climates predicted by global climate models. Despite genetic variance for traits under selection, among-trait genetic correlations that are antagonistic to the direction of selection limit adaptive evolution within these populations. Predicted rates of evolutionary response are much slower than the predicted rate of climate change.