Population genetic structure of three species in the genus Astrocaryum G. Mey. (Arecaceae).

We assessed the level and distribution of genetic diversity in three species of the economically important palm genus Astrocaryum located in Pará State, in northern Brazil. Samples were collected in three municipalities for Astrocaryum aculeatum: Belterra, Santarém, and Terra Santa; and in two municipalities for both A. murumuru: Belém and Santo Antônio do Tauá and A. paramaca: Belém and Ananindeua. Eight microsatellite loci amplified well and were used for genetic analysis. The mean number of alleles per locus for A. aculeatum, A. murumuru, and A. paramaca were 2.33, 2.38, and 2.06, respectively. Genetic diversity was similar for the three species, ranging from HE = 0.222 in A. aculeatum to HE = 0.254 in A. murumuru. Both FST and AMOVA showed that most of the genetic variation was found within populations for all three species, but high genetic differentiation among populations was found for A. aculeatum. Three loci were not in Hardy-Weinberg equilibrium, with populations of A. paramaca showing a tendency for the excess of heterozygotes (FIS = -0.144). Gene flow was high for populations of A. paramaca (Nm = 19.35). Our results suggest that the genetic diversity within populations followed the genetic differentiation among populations due to high gene flow among the population. Greater geographic distances among the three collection sites for A. aculeatum likely hampered gene flow for this species.

Limited genomic consequences of mixed mating in the recently derived sister species pair, Collinsia concolor and Collinsia parryi.

Highly selfing species often show reduced effective population sizes and reduced selection efficacy. Whether mixed mating species, which produce both self and outcross progeny, show similar patterns of diversity and selection remains less clear. Examination of patterns of molecular evolution and levels of diversity in species with mixed mating systems can be particularly useful for investigating the relative importance of linked selection and demographic effects on diversity and the efficacy of selection, as the effects of linked selection should be minimal in mixed mating populations, although severe bottlenecks tied to founder events could still be frequent. To begin to address this gap, we assembled and analysed the transcriptomes of individuals from a recently diverged mixed mating sister species pair in the self-compatible genus, Collinsia. The de novo assembly of 52 and 37 Mbp C. concolor and C. parryi transcriptomes resulted in ~40 000 and ~55 000 contigs, respectively, both with an average contig size ~945. We observed a high ratio of shared polymorphisms to fixed differences in the species pair and minimal differences between species in the ratio of synonymous to replacement substitutions or codon usage bias implying comparable effective population sizes throughout species divergence. Our results suggest that differences in effective population size and selection efficacy in mixed mating taxa shortly after their divergence may be minimal and are likely influenced by fluctuating mating systems and population sizes.

The genetic consequences of fluctuating inbreeding depression and the evolution of plant selfing rates.

The magnitude of inbreeding depression, a central parameter in the evolution of plant mating systems, can vary depending on environmental conditions. However, the underlying genetic mechanisms causing environmental fluctuations in inbreeding depression, and the consequences of this variation for the evolution of self-fertilization, have been little studied. Here, we consider temporal fluctuations of the selection coefficient in an explicit genetic model of inbreeding depression. We show that substantial variance in inbreeding depression can be generated at equilibrium by fluctuating selection, although the simulated variance tends to be lower than has been measured in experimental studies. Our simulations also reveal that purging of deleterious mutations does not depend on the variance in their selection coefficient. Finally, an evolutionary analysis shows that, in contrast to previous theoretical approaches, intermediate selfing rates are never evolutionarily stable when the variation in inbreeding depression is due to fluctuations in the selection coefficient on deleterious mutations.

Variation and constraint in plant evolution and development.

The goal of this short review is to consider the interrelated phenomena of phenotypic variation and genetic constraint with respect to plant diversity. The unique aspects of plants, including sessile habit, modular growth and diverse developmental programs expressed at the phytomer level, merit a specific examination of the genetic basis of their phenotypic variation, and how they experience and escape genetic constraint. Numerous QTL studies with wild and domesticated plants reveal that most phenotypic traits are polygenic but vary in the number and effect of the loci contributing, from a few loci of large effects to many with small effects. Further, somatic mutations, developmental plasticity and epigenetic variation, especially gene methylation, can contribute to increases in phenotypic variation. The flip side of these processes, genetic constraint, can similarly be the result of many factors, including pleiotropy, canalization and genetic redundancy. Genetic constraint is not only a mechanism to prevent change, however, it can also serve to direct evolution along certain paths. Ultimately, genetic constraint often comes full circle and is released through events such as hybridization, genome duplication and epigenetic remodeling. We are just beginning to understand how these processes can operate simultaneously during the evolution of ecologically important traits in plants.

Spatial population genetic structure in Trillium grandiflorum: the roles of dispersal, mating, history, and selection.

The roles of the various potential ecological and evolutionary causes of spatial population genetic structure (SPGS) cannot in general be inferred from the extant structure alone. However, a stage-specific analysis can provide clues as to the causes of SPGS. We conducted a stage-specific SPGS analysis of a mapped population of about 2000 Trillium grandiflorum (Liliaceae), a long-lived perennial herb. We compared SPGS for juvenile (J), nonreproductive (NR), and reproductive (R) stages. Fisher's exact test showed that genotypes had Hardy-Weinberg frequencies at all loci and stage classes. Allele frequencies did not differ between stages. Bootstrapped 99% confidence intervals (99%CI) indicate that F-statistic values are indistinguishable from zero, (except for a slightly negative FIT for the R stage). Spatial autocorrelation was used to calculate f the average kinship coefficient between individuals within distance intervals. Null hypothesis 99%CIs for f were constructed by repeatedly randomizing genotypic locations. Significant positive fine-scale genetic structure was detected in the R and NR stages, but not in the J stage. This structure was most pronounced in the R stage, and declined by about half in each remaining stage: near-neighbor f = 0.122, 0.065, 0.027, for R, NR, and J, respectively. For R and NR, the near-neighbor f lies outside the null hypothesis 99%CI, indicating kinship at approximately the level of half-sibs and first cousins, respectively. We also simulated the expected SPGS of juveniles post dispersal, based on measured R-stage SPGS, the mating system, and measured pollen and seed dispersal properties. This provides a null hypothesis expectation (as a 99%CI) for the J-stage correlogram, against which to test the likelihood that post-dispersal events have influenced J-stage SPGS. The actual J correlogram lies within the null hypothesis 99%CI for the shortest distance interval and nearly all other distance intervals indicating that the observed low recruitment, random mating and seed dispersal patterns are sufficient to account for the disappearance of SPSG between the R and the J stages. The observed increase in SPGS between J and R stages has two potential explanations: history and local selection. The observed low total allelic diversity is consistent with a past bottleneck: a possible historical explanation. Only a longitudinal stage-specific study of SPGS structure can distinguish between historical events and local selection as causes of increased structure with increasing life history stage.

Family- and population-level responses to atmospheric CO2 concentration: gas exchange and the allocation of C, N, and biomass in Plantago lanceolata (Plantaginaceae).

To ascertain the inheritance of responses to changing atmospheric CO(2) content, we partitioned response to elevated CO(2) in Plantago lanceolata between families and populations in 18 families in two populations. Plants were grown in 35 Pa and 71 Pa partial pressure of CO(2) (pCO(2)) in open-top chambers. We measured above- and belowground mass, carbon (C), nitrogen (N), hexose sugar, and gas exchange properties in both CO(2) treatments. Families within populations differed in mass, mass allocation, root : shoot ratios, aboveground percentage N, C : N ratio, and gas exchange properties. The CO(2) × family interaction is the main indicator of potential evolutionary responses to changing CO(2). Significant CO(2) × family interactions were observed for N content, C : N ratio, and photosynthetic rate (A: instantaneous light-saturated carbon assimilation capacity), intercellular CO(2) concentration, transpiration rate (E), and water use efficiency (WUE = A/E), but not for stomatal conductance. Families differed significantly in acclimation across time. The ratio of A in elevated vs. ambient growth CO(2), when measured at a common internal CO(2) partial pressure was 0.79, indicating down-regulation of A under CO(2) enrichment. Mass, C : N ratio, percentage, C (%C), and soluble sugar all increased significantly but overall %N did not change. Increases in %C and sugar were significant and were coincident with redistribution of N aboveground. The observed variation among populations and families in response to CO(2) is evidence of genetic variation in response and therefore of the potential for novel evolutionary trajectories with rising atmospheric CO(2).

Sex among the flowers: the distribution of plant mating systems.

Previous reviews of plant outcrossing rate survey data have agreed that predominant selfing and predominant outcrossing are alternative stable states of mating system evolution. We reanalyzed the most recent data and plot outcrossing rates as a continuous variable rather than as a class variable. Wind-pollinated species are indeed bimodal. However, the shape of the distributions for animal-pollinated species reveals that intermediate rates of outcrossing are common (49% of species fall between 20% and 80% outcrossing). Consequently, we suggest that mating system is best considered a continuous rather than a discrete character of plant populations.

The mechanism of delayed selfing in Collinsia verna (Scrophulariaceae).

Collinsia verna, blue-eyed Mary, has floral attributes of an outcrossing species, yet most flowers readily self-pollinate under greenhouse conditions. Here we describe the mechanism of self-pollination in C. verna via changes in relative positions of the stigma and anthers and late timing of receptivity, resulting in delayed selfing. Each flower contains four anthers that dehisce sequentially over ∼1 wk. Pollen that is not collected by pollinators accumulates in the keel petal and retains high viability (>80% pollen germination) up to the time of corolla abscission. The stigmatic surface does not become receptive until after the third anther dehisces. This overlap in the sexual phases is concurrent with a change in herkogamy during floral development. In most flowers (70%), the stigma has moved to the front of the keel and is positioned near the anthers when the third anther dehisces. Under field conditions, fruiting success of plants within pollinator exclosures was ∼75% of the fruiting success in open-pollinated plants (33% fruiting success via autogamy vs. 44% fruiting success, respectively). Collinsia verna plants in pollinator exclosures exhibit variation in autogamy rates within natural populations (range 0-80%). In addition, only half of naturally pollinated, receptive flowers examined had pollen tubes growing in their styles. In contrast, shortly after corolla abscission, nearly all flowers examined (96%) had pollen tubes in their styles. Thus we find that in C. verna, autogamy occurs late in floral development, which has the potential to provide substantial reproductive assurance, and that individuals vary in their ability to set fruit through this mechanism. We suggest that delayed selfing mechanisms may be overlooked in other species and that variable pollinator availability may play a significant role in the maintenance of mixed mating in species with delayed selfing, such as C. verna.