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Hough, Josh; Immler, Simone; Barrett, Spencer C. H.; Otto, Sarah P. 2013-01-16 Frequency-dependent selection should drive dioecious populations toward a 1:1 sex ratio, but biased sex ratios are widespread, especially among plants with sex chromosomes. Here, we develop population genetic models to investigate the relationships between evolutionarily stable sex ratios, haploid selection, and deleterious mutation load. We confirm that when haploid selection acts only on the relative fitness of X and Y-bearing pollen and the sex ratio is controlled by the maternal genotype, seed sex ratios evolve toward 1:1. When we also consider haploid selection acting on deleterious mutations, however, we find that biased sex ratios can be stably maintained, reflecting a balance between the advantages of purging deleterious mutations via haploid selection, and the disadvantages of haploid selection on the sex ratio. Our results provide a plausible evolutionary explanation for biased sex ratios in dioecious plants, given the extensive gene expression that occurs across plant genomes at the haploid stage.
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Yakimowski, Sarah B.; Barrett, Spencer C. H. 2014-06-03 Clonality is often implicated in models of the evolution of dioecy, but few studies have explicitly compared clonal structure between plant sexual systems, or between the sexes in dioecious populations. Here, we exploit the occurrence of monoecy and dioecy in clonal Sagittaria latifola (Alismataceae) to evaluate two main hypotheses: (1) clone sizes are smaller in monoecious than dioecious populations, because of constraints imposed on clone size by costs associated with geitonogamy; (2) in dioecious populations, male clones are larger and flower more often than female clones because of sex-differential reproductive costs. Differences in clone size and flowering could result in discordance between ramet- and genet-based sex ratios. We used spatially explicit sampling to address these hypotheses in 10 monoecious and 11 dioecious populations of S. latifolia at the northern range limit in eastern N. America. In contrast to our predictions, monoecious clones were significantly larger than dioecious clones, probably due to their higher rates of vegetative growth and corm production, and in dioecious populations there was no difference in clone size between females and males; ramet- and genet-based sex ratios were therefore highly correlated. Genotypic diversity declined with latitude for both sexual systems, but monoecious populations exhibited lower genotypic richness. Differences in life history between the sexual systems of S. latifolia appear to be the most important determinants of clonal structure and diversity.
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Timerman, David; Barrett, Spencer C.H.; Barrett, Spencer C. H. 2018-12-11 Wind pollination has evolved from insect pollination in numerous angiosperm lineages and is associated with a characteristic syndrome of morphological traits. The traits initiating transitions to wind pollination and the ecological drivers involved are poorly understood. Here, we examine this problem in Thalictrum pubescens, an ambophilous (insect and wind pollination) species that probably represents a transitional state in the evolution of wind pollination in some taxa. We investigated wind-induced pollen release by forced harmonic motion by measuring stamen natural frequency (fn), a key vibration parameter, and its variability among nine populations. We assessed the repeatability of fn over consecutive growing seasons, the effect of this parameter on pollen release in a wind tunnel, and male reproductive success in the field using experimental manipulation of the presence or absence of pollinators. We found significant differences among populations and high repeatability within genotypes in fn. The wind tunnel assay revealed a strong negative correlation between fn and pollen release. Siring success was greatest for plants with lower fn when pollinators were absent; but this advantage diminished when pollinators were present. Our biomechanical analysis of the wind-flower interface has identified fn as a key trait for understanding early stages in the transition from insect to wind pollination.
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Sandler, George; Beaudry, Felix E.G.; Barrett, Spencer C.H.; Wright, Stephen I.; Beaudry, Felix E. G.; Barrett, Spencer C. H. 2018-05-21 The evolution of sex chromosomes is usually considered to be driven by sexually antagonistic selection in the diploid phase. However, selection during the haploid gametic phase of the lifecycle has recently received theoretical attention as possibly playing a central role in sex chromosome evolution, especially in plants where gene expression in the haploid phase is extensive. In particular, male-specific haploid selection might favour the linkage of pollen beneficial alleles to male sex determining regions on incipient Y chromosomes. This linkage might then allow such alleles to further specialise for the haploid phase. Purifying haploid selection is also expected to slow the degeneration of Y-linked genes expressed in the haploid phase. Here, we examine the evolution of gene expression in flower buds and pollen of two species of Rumex to test for signatures of haploid selection acting during plant sex chromosome evolution. We find that genes with high ancestral pollen expression bias occur more often on sex chromosomes than autosomes and that genes on the Y chromosome are more likely to become enriched for pollen expression bias. We also find that genes with low expression in pollen are more likely to be lost from the Y chromosome. Our results suggest that sex-specific haploid selection during the gametophytic stage of the lifecycle may be a major contributor to several features of plant sex chromosome evolution.
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Timerman, David; Barrett, Spencer C. H.; Barrett, Spencer C.H. 2019-01-17 1. In clonal dioecious plants, the frequency and spatial distribution of flowering ramets contains information on the underlying genet sex ratio. These measures can also provide insight on potential ecological mechanisms causing variation and bias in sex ratios among populations. 2. We used a novel likelihood-based approach and spatial clustering model to estimate the genet sex ratios from flowering ramet data collected from 32 populations of dioecious Thalictrum pubescens, a clonal species from eastern N. America that occupies moist wetland and forested environments. We investigated sex ratios of seed families, clone size, patterns of flowering and plant height to determine potential causes of sex ratio bias. 3. Flowering ramet sex ratios varied considerably among populations but were significantly male-biased. Seed families grown to flowering also exhibited the same degree of male bias. Both models predicted close correspondence between ramet and genet sex ratios. The likelihood model revealed that gender differences in ramet production could not account for biased sex ratios. The spatial clustering model indicated that ramets were significantly clustered at two spatial scales and estimated similar cluster sizes and densities for both sexes. There was no evidence for spatial segregation of the sexes. Both sexes were equally likely to flower in consecutive years and repeated bouts of flowering had no effect on ramet height. 4. Synthesis. Our analyses suggest that the widespread occurrence of male-biased sex ratios in T. pubescens is unlikely to result from sexual differences in clonal growth or habitat preferences. The bias appears to become established early in the life cycle, perhaps at the seed stage as consequence of local resource competition.
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Percy, Diana M.; Argus, George W.; Cronk, Quentin C.; Fazekas, Aron J.; Kesanakurti, Prasad R.; Burgess, Kevin S.; Husband, Brian C.; Newmaster, Steven G.; Barrett, Spencer C. H.; Graham, Sean W.; Barrett, Spencer C.H. 2014-06-17 Willows (Salix: Salicaceae) form a major ecological component of Holarctic floras, and consequently are an obvious target for a DNA-based identification system. We surveyed two to seven plastid genome regions (~3.8 kb; ~3% of the genome) from 71 Salix species across all five subgenera, to assess their performance as DNA barcode markers. Although Salix has a relatively high level of interspecific hybridization, this may not sufficiently explain the near complete failure of barcoding that we observed: only one species had a unique barcode. We recovered 39 unique haplotypes, from more than 500 specimens, that could be partitioned into six major haplotype groups. A unique variant of group I (haplotype 1*) was shared by 53 species in three of five Salix subgenera. This unusual pattern of haplotype sharing across infrageneric taxa is suggestive of either a massive non-random coalescence failure (incomplete lineage sorting), or of repeated plastid capture events, possibly including a historical selective sweep of haplotype 1* across taxonomic sections. The former is unlikely as molecular dating indicates that haplotype 1* originated recently, and is nested in the oldest major haplotype group in the genus. Further, we detected significant non-neutrality in the frequency spectrum of mutations in group I, but not outside group I, and demonstrated a striking absence of geographic structure to the haplotype distributions in this group. The most likely explanation for the patterns we observed involves recent repeated plastid capture events, aided by widespread hybridization and long-range seed dispersal, but primarily propelled by one or more trans-species selective sweeps.

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