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The life cycles of sexually reproducing animals and flowering plants begin with male and female gametes and their fusion to form a zygote. Selection at this earliest stage is crucial for offspring quality and raises similar evolutionary issues, yet zoology and botany use dissimilar approaches. There are striking parallels in the role of prezygotic competition for sexual selection on males, cryptic female choice, sexual conflict, and against selfish genetic elements and genetic incompatibility. In both groups, understanding the evolution of sex-specific and reproductive traits will require an appreciation of the effects of prezygotic competition on fitness., Multicellular animals and flowering plants (angiosperms) spend most of their life cycle in the diploid state. This cycle, however, begins with a short haploid phase, which ends when male and [...]

A study describes the capacity for floral phenotype to respond to selection through a review of published data on heritabilities and genetic correlations for several classes of floral traits in hermaphroditic plants. Among the findings is a significant heritability for all floral traits, with variation among them, as well as a tendency for heritability to vary with mating system, but not life history.

Abstract: Since the launch of Ecology Letters in 1998, the field of Pollination Ecology has changed considerably in its focus. In this review, we discuss the major discoveries across the past two decades. We quantitatively synthesise the frequency by which different concepts and topics appeared in the peer‐reviewed literature, as well as the connections between these topics. We then look forward to identify pressing research frontiers and opportunities for additional integration in the future. We find that there has been a shift towards viewing plant–pollinator interactions as networks and towards understanding how global drivers influence the plants, pollinators and the ecosystem service of pollination. Future frontiers include moving towards a macroecological view of plant–pollinator interactions, understanding how ecological intensification and urbanisation will influence pollination, considering other interactions, such as plant–microbe–pollinator networks, and understanding the causes and consequences of extinctions. Pollination Ecology is poised to advance our basic understanding of the ecological and evolutionary factors that shape plant–animal interactions and to create applied knowledge that informs conservation decision making. [ABSTRACT FROM AUTHOR]

Sexual selection can occur in plants even though individuals do not directly interact via a process known as interaction-independent sexual selection. For example, if variation exists among individuals in flower production and this affects their attractiveness to pollinators, then individuals with relatively high flower production may achieve relatively high mating success via pollen donation. We review research on a variety of traits in flowering plants that affect pollinator visitation, with the underlying premise being that these can influence mate acquisition. In some cases research has shown that phenotypes that enhance pollen donation can be antagonistic to fitness achieved via female function, while in other cases both sex functions select for the same phenotype. We also highlight different approaches researchers have taken to address these issues, including studies on gynodioecious species, which contain both hermaphrodites and females. Examples include manipulations of flowers to hone in on the traits that actually contribute to pollinator discrimination, phenotypic selection analyses, and quantitative-genetic parameter estimation.

The evolutionary transition from hermaphroditism (combined sexes) to dioecy (separate sexes) is associated with whole genome duplication (polyploidy) in several flowering plant genera. Moreover, there is evidence for transitions in the opposite direction, i.e. a loss of dioecy with an increase in ploidy. Here, we review evidence for these associations, synthesize previous ideas on the mechanism underlying the patterns and explore alternative pathways. Specifically, we examine potential ecological and genetic mechanisms, differentiated by whether ploidy or gender (functional sex expression of the plant) changes are the primary cause and whether the effect is direct or indirect. An analysis of 22 genera variable for both ploidy and gender indicates that gender monomorphism (hermaphroditism, monoecy) is more common among diploid than polyploid species, whereas gender dimorphism (dioecy, gynodioecy, subdioecy) is more frequent among polyploid species. The transition from diploid hermaphroditic to polyploid gender-dimorphic taxa may arise directly through changes in gender as a result of genome duplication through genomic rearrangements or homeologous recombination, or changes in gender may result in increased unreduced gamete production leading to polyploid formation. Alternatively, the transition may occur through the indirect effects of genome duplication on mating system and inbreeding depression, which favor selection for unisexuality, or habitat shifts associated with unisexuality may simultaneously cause increased unreduced gamete production. Novel mechanisms for transitions in the opposite direction (from dioecy to hermaphroditism with increase in ploidy) include disruption of genetic sex determination and the benefits of reproductive assurance. We highlight key questions requiring further attention and promising approaches for answering them and better clarifying the genesis of sexual system polyploidy associations. See also the sister article focusing on animals by Wertheim et al. in this themed issue. Copyright © 2013 S. Karger AG, Basel [ABSTRACT FROM AUTHOR]

We studied the natural hybrid (Fragaria × ananassa subsp. cuneifolia) between two sexually dimorphic octoploid strawberry species (Fragaria virginiana and Fragaria chiloensis) to gain insight into the dynamics of sex chromosomes and the genesis of sexual dimorphism. Male sterility is dominant in both the parental species and thus will be inherited maternally, but the chromosome that houses the sex-determining region differs. Thus, we asked whether (1) the cytotypic composition of hybrid populations represents one or both maternal species, (2) the sex-determining chromosome of the hybrid reflects the location of male sterility within the maternal donor species and (3) crosses from the hybrid species show less sexual dimorphism than the parental species. We found that F. × ananassa subsp. cuneifolia populations consisted of both parental cytotypes but one predominated within each population. Genetic linkage mapping of two crosses showed dominance of male sterility similar to the parental species, however, the map location of male sterility reflected the maternal donor in one cross, but not the other. Moreover, female function mapped to a single region in the first cross, but to two regions in the second cross. Aside from components of female function (fruit set and seed set), other traits that have been found to be significantly sexually dimorphic in the pure species were either not dimorphic or were dimorphic in the opposite direction to the parental species. These results suggest that hybrids experience some disruption of dimorphism in secondary sexual traits, as well as novel location and number of quantitative trait locus (QTL) affecting sex function. [ABSTRACT FROM AUTHOR]

Florivores are antagonists that damage flowers, and have direct negative effects on flowering and pollination of the attacked plants. While florivory has mainly been studied for its consequences on seed production or siring success, little is known about its impact on mating systems. Damage to flowers can alter pollinator attraction to the plant and may therefore modify patterns of pollen transfer. However, the consequences of damage for mating systems can take two forms: a decrease in flower number reduces opportunities for intra-inflorescence pollen deposition (geitonogamy), which, in turn, may lead to a decrease in selfing; whereas a decrease in floral display may also reduce overall visitation and thus increase the chances of self-pollination via facilitated or autonomous autogamy. We investigated the effects of damage by a bud-clipping weevil ( Anthonomus signatus) in Fragaria virginiana in an experimental setting mimicking natural conditions. We found that increased damage led to an increase in selfing, a result consistent with the increased autogamy pathway. We discuss the implications of this finding and evaluate the generality of florivore-mediated mating system expression. [ABSTRACT FROM AUTHOR]

The evolution of separate sexes (dioecy) from hermaphroditism is one of the major evolutionary transitions in plants, and this transition can be accompanied by the development of sex chromosomes. Studies in species with intermediate sexual systems are providing unprecedented insight into the initial stages of sex chromosome evolution. Here, we describe the genetic mechanism of sex determination in the octoploid, subdioecious wild strawberry, Fragaria virginiana Mill., based on a whole-genome simple sequence repeat (SSR)-based genetic map and on mapping sex determination as two qualitative traits, male and female function. The resultant total map length is 2373 cM and includes 212 markers on 42 linkage groups (mean marker spacing: 14 cM). We estimated that approximately 70 and 90% of the total F. virginiana genetic map resides within 10 and 20 cM of a marker on this map, respectively. Both sex expression traits mapped to the same linkage group, separated by approximately 6 cM, along with two SSR markers. Together, our phenotypic and genetic mapping results support a model of gender determination in subdioecious F. virginiana with at least two linked loci (or gene regions) with major effects. Reconstruction of parental genotypes at these loci reveals that both female and hermaphrodite heterogamety exist in this species. Evidence of recombination between the sex-determining loci, an important hallmark of incipient sex chromosomes, suggest that F. virginiana is an example of the youngest sex chromosome in plants and thus a novel model system for the study of sex chromosome evolution.Heredity (2008) 101, 507–517; doi:10.1038/hdy.2008.100; published online 17 September 2008 [ABSTRACT FROM AUTHOR]

The hypothesis that flower maintenance requires resources that wouldbe used to support other plant functions (i.e. a cost of floral maintenance) was tested by experimentally manipulating floral longevity. Plants of Clarkia tembloriensis, a species with pollination-induced flower senescence, received either early or late pollinations (long and short longevities, respectively). We examined the effect of this manipulation on (1) perflower allocation to nectar production and (2) flower, fruit and seed production per plant under two levels of resource availability. The direct costs of floral longevity measured in terms of nectar sugar were high: flowers that were maintained 35% longerinvested proportionately more in nectar sugar (30%). At the whole-plant level, a cost of floral longevity was manifested as reduced seed production, but the magnitude of this cost varied with resource level. While plants with longer-lived flowers showed a 12% reduction in seed production, those that experienced reduced resource levels via partial defoliation, showed a decrement in seed production that was almost three times larger (34%). These differences were not brought aboutby changes in the number of flowers and fruits, but by significant alterations in their sizes. A model that expresses the cost of flower maintenance as a trade-off between floral longevity and seed production shows that an optimal flower longevity is determined by both the rate of fitness accrual and the cost of floral maintenance. [ABSTRACT FROM AUTHOR]

Plant reproduction relies on transfer of pollen from anthers to stigmas, and the majority of flowering plants depend on biotic or abiotic agents for this transfer. A key metric for characterizing if pollen receipt is insufficient for reproduction is pollen limitation, which is assessed by pollen supplementation experiments. In a pollen supplementation experiment, fruit or seed production by flowers exposed to natural pollination is compared to that following hand pollination either by pollen supplementation (i.e. manual outcross pollen addition without bagging) or manual outcrossing of bagged flowers, which excludes natural pollination. The GloPL database brings together data from 2969 unique pollen supplementation experiments reported in 927 publications published from 1981 to 2015, allowing assessment of the strength and variability of pollen limitation in 1265 wild plant species across all biomes and geographic regions globally. The GloPL database will be updated and curated with the aim of enabling the continued study of pollen limitation in natural ecosystems and highlighting significant gaps in our understanding of pollen limitation.

We explored the idea that sex ratio represents a unique context for selection on attractive traits by manipulating sex ratio and pollinator abundance in experimental populations of a gender-dimorphic wild strawberry Fragaria virginiana. We found that increasing the frequency of functional males (the pollen-bearing morph) increased the frequency of pollen-collecting syrphid flies in the pollinator assemblage, decreased pollinator visitation to less preferred morph (females), and decreased the degree of pollen limitation of females. Moreover, sex ratio influenced the strength of selection on petal size through female fitness but did not alter the strength of selection through male fitness components, suggesting that sex ratio can alter the gender bias of selection on an attractive trait. This study of context-dependent selection has important implications for the evolution of sexual dimorphism in attractive traits. First, it suggests that only certain conditions generate male-biased selection and, thus, could lead to selection-driven male-biased petal size dimorphism. Second, it suggests that flexible pollinator foraging may be an important mechanism by which sex ratio influences selection on attractive traits. Finally, it implies that variation in sex ratio could limit the evolution of sexual dimorphism and/or could maintain genetic variation in attractive traits.

In this study we dissect the causes of variation in intra-inflorescence allocation in a sexually polymorphic species, Fragaria virginiana. We separated out the effects of resource competition during flowering from those of inflorescence architecture, as well as identified the effects of sex morph and genotype. We found position-based variation in petal length, ovule, pollen, and flower number to be influenced more by architecture than by our resource manipulations during flowering. We also found both genotype- and sex-specific intra-inflorescence patterns. Furthermore, our data indicate that the sex morph-specific intra-inflorescence patterns result from architectural modifications of the basic pattern. In fact, sex-differential intra-inflorescence patterns suggest that fitness through male and female function may be maximized by different resource distribution patterns within the inflorescence and may have been modified by past selection. Specifically, females invested heavily in ovules at positions where fruit set was most likely (primary and secondary), at the expense of flower number and allocation per flower at more distal positions. Whereas functional males invested minimally in ovules at all flower positions and produced the most abundantly flowered inflorescences, hermaphrodites, on the other hand, showed intermediate patterns, implying a compromise between sex functions. We suggest that consideration of intra-inflorescence allocation and inflorescence architecture may reveal the mechanism underlying sexual dimorphism in flower allocation and number.

The independent evolution of floral phenotype is an important part of the process of gender specialization during the evolution of dioecy from hermaphroditism. However, we have little information on the genetic variation of floral traits in species with separate genders. Gynodioecious species (co-occurrence of females and hermaphrodites) have a breeding system intermediate between hermaphroditism and complete separation of the sexes (dioecy) and thus can provide insight into the genetic architecture underlying floral phenotype with respect to both primary (stamens and carpels) and secondary (petals) sexual traits. I used a nested breeding design to examine the potential for response to selection on floral traits and to examine whether this response would be similar in the two sex morphs of gynodioecious Fragaria virginiana. There was significant genetic variation underlying all floral traits, although narrow-sense heritabilities (ranging from -0.25 to 0.44) were, in most cases, much lower than broad-sense ones (ranging from 0.28 to 1. 53). Moreover, the sex morphs differed significantly in their heritabilities for shared traits, such as stamen length, and showed a tendency towards differing significantly in others, like carpel number and petal length. In addition, correlations between the sex morphs for these traits (ranging from 0.41 to 0.58) were significantly greater than 0, but less than 1. These results indicate that greater sexual dimorphism could evolve in this population of F. virginiana, even if selection on these traits is not divergent. However, strong developmental integration of floral traits (e.g. stamen length and petal length) and high levels of nonadditive genetic variance may represent barriers to the evolution of complete sexual dimorphism.