How geitonogamous selfing affects sex allocation in hermaphrodite plants

Does the mode of self-pollination affect the evolutionarily stable allocation to male vs. female function? We distinguish the following scenarios. (1) An ‘autogamous’ species, in which selfing occurs within the flower prior to opening. The pollen used in selfing is a constant fraction of all pollen grains produced. (2) A species with ‘abiotic pollination’, in which selfing occurs when pollen dispersed in one flower lands on the stigma of a nearby flower on the same plant (geitonogamy). The selfing rate increases with male allocation but a higher selfing rate does not mean a reduced export of pollen. (3) An ‘animal-pollinated’ species with geitonogamous selfing. Here the selfing rate also increases with male allocation, but pollen export to other plants in the population is a decelerating function of the number of simultaneously open flowers. In all three models selfing selects for increased female allocation. For model 3 this contradicts the general opinion that geitonogamous selfing does not affect evolutionarily stable allocations. In all models, the parent benefits more from a female-biased allocation than any other individual in the population. In addition, in models 2 and 3, greater male allocation results in more local mate competition. In model 3 and in model 2 with low levels of inbreeding depression, hermaphroditism is evolutionarily stable. In model 2 with high inbreeding depression, the population converges to a fitness minimum for the relative allocation to male function. In this case the fitness set is bowed inwards, corresponding with accelerating fitness gain curves. If the selfing rate increases with plant size, this is a sufficient condition for size-dependent sex allocation (more allocation towards seeds in large plants) to evolve. We discuss our results in relation to size-dependent sex allocation in plants and in relation to the evolution of dioecy.     

Evidence for maintenance of sex by pathogens in plants

The predominance of outcrossing despite the substantial transmission advantage of self-fertilization remains a paradox. Theory suggests that selection can favor outcrossing if it enables the production of offspring that are less susceptible to pathogen attack than offspring produced via self-fertilization. Thus, if pathogen pressure is contributing to the maintenance of outcrossing in plants, there may be a positive correlation between the number of pathogen species attacking plant species and the outcrossing rate of the plant species. We tested this hypothesis by examining the association between outcrossing rate and the number of fungal pathogen species that attack a large, taxonomically diverse set of seed plants. We show that plant species attacked by more fungal pathogen species have higher outcrossing rates than plants with fewer enemies. This relationship persists after correcting for study bias among natural and agricultural species of plants. We also accounted for the nested hierarchy of relationships among plant lineages by conducting phylogenetically independent contrasts (PICs) within genera and families that were adequately represented in our dataset. A meta-analysis of the correlation between pathogen and outcrossing PICs shows that there is a positive correlation between pathogen species number and outcrossing rates. This pattern is consistent with the hypothesis that pathogen-mediated selection may contribute to the maintenance of outcrossing in species of seed plants.     

Optimal allocation to vegetative and sexual reproduction in plants: the effect of ramet density

Vegetative reproduction is a very common alternative by which plants can contribute to the next generations. There are many considerations predicting which mode of reproduction, vegetative or sexual, should be favored and numerous experimental studies to verify them. However, the results are inconsistent especially when the effect of plant density is considered. I apply here a dynamic optimization model to predict the rate of vegetative and sexual reproduction in plants as a response to changes in the local plant density. The population is assumed to occupy a heterogeneous environment consisting of patches in which growth and reproduction of plants are possible and unfavorable space between them. As the environment is globally stable, the seeds, which can disperse without restriction, exhibit a constant recruitment rate. The ramets are assumed to settle only within the patch of the mother plant. The rate of ramet production effects local density, which in turn determines ramet recruitment. The optimal strategy maximizes the expected lifetime genetic contribution, realized via both vegetative and sexual reproduction. The solutions obtained under these assumptions are dualistic. The model predicts that different approaches applied in studying the effect of ramet density should give opposite outcomes. When the comparison is between patches in natural populations, a positive relationship between relative ramet allocation and density is expected. When the density is experimentally manipulated or its effect is analyzed across different successional stages, a negative relationship should be found. The results seem to be confirmed by empirical studies.     

The loss of sex in clonal plants

Most plants combine sexual and clonal reproduction, and the balance between the two may vary widely between and within species. There are many anecdotal reports of plants that appear to have abandoned sex for clonal reproduction, yet few studies have quantified the degree of sexual variation in clonal plants and fewer still have determined the underlying ecological and/or genetic factors. Recent empirical work has shown that some clonal plants exhibit very wide variation in sexual reproduction that translates into striking variation in genotypic diversity and differentiation of natural populations. Reduced sexual reproduction may be particularly common at the geographical margins of species' ranges. Although seed production and sexual recruitment may often be limited by biotic and abiotic aspects of the environment in marginal populations, genetic factors, including changes in ploidy and sterility mutations, may also play a significant role in causing reduced sexual fertility. Moreover, environmental suppression of sexual recruitment may facilitate the evolution of genetic sterility because natural selection no longer strongly maintains the many traits involved in sex. In addition to the accumulation of neutral sterility mutations in highly clonal populations, the evolution of genetic infertility may be facilitated if sterility is associated with enhanced vegetative growth, clonal propagation or survival through either resource reallocation or pleiotropy. However, there are almost no experimental data with which to distinguish among these possibilities. Ultimately, wide variation in genotypic diversity and gene flow associated with the loss of sex may constrain local adaptation and the evolution of the geographical range limit in clonal plants.     

雌雄同花植物的性分配

性分配理论假定雌雄功能之间存在着trade-off,对一种性别的投入增多必然会减少对另一性别的投入。雌雄功能投入的适合度曲线的形状决定了哪种繁育系统是进化稳定的。因此,性分配理论可以解释植物繁育系统的进化,尤其被认为是雌雄异株进化的选择机制之一。目前的实验研究分别在物种间、种群间、个体间及花间四个层次上进行:自交率的程度对物种和种群的性分配都有影响;虫媒和风媒植物的性分配是个体大小依赖的;而且花序内花的性分配模式受昆虫访花行为的影响。相对于理论,性分配的实验研究明显滞后,随着分子标记技术的普及,性分配理论将会获得更大的发展。繁殖分配需要进一步与性分配理论结合,尤其在空间尺度上资源分配与繁育系统变化的研究是很有意义的。     

植物生长发育中程序性细胞死亡

本文简要介绍了植物细胞凋亡的一些特点以及植物在营养生长和生殖生长过程中发生的细胞凋亡现象。指出细胞凋亡是植物生长发育过程中正常的生理现象。     

Size and sex allocation in monoecious woody plants

The female size advantage hypothesis predicts that the allocation ratio of female: male reproductive effort should increase with plant size (total reproductive effort). A male height advantage hypothesis has also been proposed, based on the supposed greater advantage of height to male reproductive success in wind-pollinated plants. These ideas were tested with data for wind-pollinated, monoecious trees and shrubs which exhibit a suitably large range of sizes. Number of male inflorescences increased faster with size than did number of female inflorescences in 2 of 9 species; in the remaining 7 species there was no significant difference. The male:female ratio of inflorescence numbers increased with height in 4 of 7 species and did not change significantly in the remaining 3 species, as shown by regression. Height and size are highly correlated and so their effects could not be distinguished. The fact that many conifers place the female cones uppermost in the crown suggests that size and not height favors increased allocation to male function, as does well-established theory connecting the existence of male versus female size advantage to pollen and seed dispersal chacteristics. Regression analysis of the relation between male and female reproductive effort should be done by reduced major axis regression; ordinary least squares regression underestimates slopes; in this study opposite conclusions could be drawn from ordinary least squares and reduced major axis regressions.     

性的进化:起源和维持

有性繁殖是真核生物生命周期的一个普遍特征。然而,在解释有性繁殖的这种普遍存在时却遇到了许多的困难,包括雄性的双重损失和重组负荷等。那么,有性繁殖是如何起源和维持的？人们提出了许多可能解释的假说,但是到目前为止还没有一个明确的答案。作者就众多的理论做一个简明综述。     

A field survey of dioecious plants in Israel: sex ratio in seven rare species

Sex ratio was tested in seven rare dioecious species in Israel. This was in order to complete a previously published sex ratio survey of the other 41 dioecious species in Israel, and to examine further the hypothesis that sex ratio in natural populations is usually 1:1. One population was tested in each of five rare wild species. In addition, one rare feral population was tested in each of two non-native, naturalized tree species. Sex ratio in all seven species tested was not different from the expected 1:1. These results strengthen the suggestion that in natural populations of dioecious plants sex ratio is usually 1:1.     

Chromosomal variability in tissue cultures and regenerated plants of Hordeum

Summary Spontaneous polyploidy, aneuploidy, and chromosomal rearrangements were observed in callus and suspension cultures of Hordeum vulgare, H. jubatum, and their interspecific hybrid. The extent to which each class of chromosomal variability was present in a culture depended upon differentiated state, age, and history. Cytological and isozymic analysis of subdivided callus cultures revealed spatial segregation of chromosomal variability. Cytogenetic analyses were performed to determine the expression of this in vitro chromosomal variability in corresponding regenerated plant tissues. A complete loss of polyploidy and a decrease in aneuploidy and chromosomal rearrangements were observed. Analyses of specific isozyme activities in regenerates suggested that a quantitative segregation of H. vulgare and H. jubatum genomes had occurred in tissue cultures of their interspecific hybrid. Possible uses of in vitro chromosomal variability for plant breeding and genetical studies are discussed.Work supported by a grant to Project 3195 from the Michigan Agricultural Experiment Station and by Grant E4-76-S-02-7528. A004 to P.S. Carlson from the US Department of Energy     

Testing the predictions of sex allocation hypotheses in dimorphic,cooperatively breeding riflemen

Evolutionary theory predicts that parents should invest equally in the two sexes. If one sex is more costly, a production bias is predicted in favour of the other. Two well‐studied causes of differential costs are size dimorphism, in which the larger sex should be more costly, and sex‐biased helping in cooperative breeders, in which the more helpful sex should be less costly because future helping “repays” some of its parents’ investment. We studied a bird species in which both processes should favor production of males. Female riflemen Acanthisitta chloris are larger than males, and we documented greater provisioning effort in more female‐biased broods indicating they are likely costlier to raise. Riflemen are also cooperative breeders, and males provide more help than females. Contrary to expectations, we observed no male bias in brood sex ratios, which did not differ significantly from parity. We tested whether the lack of a population‐wide pattern was a result of facultative sex allocation by individual females, but this hypothesis was not supported either. Our results show an absence of adaptive patterns despite a clear directional hypothesis derived from theory. This appears to be associated with a suboptimal female‐biased investment ratio. We conclude that predictions of adaptive sex allocation may falter because of mechanistic constraint, unrecognized costs and benefits, or weak selection.     

Spatial seed and pollen games: dispersal,sex allocation,and the evolution of dioecy

The evolutionary forces shaping within‐ and across‐species variation in the investment in male and female sex function are still incompletely understood. Despite earlier suggestions that in plants the evolution or cosexuality vs. dioecy, as well as sex allocation among cosexuals, is affected by seed and pollen dispersal, no formal model has explicitly used dispersal distances to address this problem. Here, we present a game‐theory model as well as a simulation study that fills in this gap. Our model predicts that dioecy should evolve if seeds and pollen disperse widely and that sex allocation among cosexuals should be biased towards whichever sex function produces more widely dispersing units. Dispersal limitations stabilize cosexuality by reinforcing competition between spatially clumped dispersal units from the same source, leading to saturating fitness returns that render sexual specialization unprofitable. However, limited pollen dispersal can also increase the risk of selfing, thus potentially selecting for dioecy as an outbreeding mechanism. Finally, we refute a recent claim that cosexuals should always invest equally in both sex functions.     

DNA repair in higher plants

Numerous studies have demonstrated a requirement in plants for repair of DNA damage arising from either intrinsic or extrinsic sources. Investigations also have revealed a capacity for repair types of DNA damage, and conversely, identified mutants apparently defective in such repair. This article provides a concise overview of nuclear DNA repair mechanisms in higher plants, particularly those processes concerned with the repair of UV-induced lesions, and includes surveys of UV-sensitive mutants and genes implicated in DNA repair.     

Senescence in growth and reproductive allocation in a bunchgrass

• Senescence is a puzzling phenomenon. Few convincing studies of senescence in perennial herbaceous plants exist. While ramets are known to senesce, whether senescence of bunchgrasses actually occurs is not clear.
• In this study, we grew a set of plants of Elymus excelsus, a bunchgrass, to examine plant size, sexual reproduction and bud formation in individual plants in relation to their gradual ageing, in order to determine whether E. excelsus experiences senescence. We collected data in two consecutive years (2009 and 2010) from field samples of plants from 1 to 5 years old. Using regression models, we performed age‐related analyses of growth and reproduction parameters.
• Our results showed that individual plant size (diameter, individual biomass), total biomass of ramets, number and biomass of reproductive ramets, percentage of ramets that were reproductive, reproductive allocation, over‐wintering buds and juvenile ramets all declined with age. However, vegetative growth (number and biomass of vegetative ramets) did not decrease with age.
• Those plants that survived, dwindled in size as they aged. However, no plants shifted their resource allocation between growth and reproduction as they aged, so the shift in allocation did not account for the fall in size.

     

Sex, outcrossing and mating types: unsolved questions in fungi and beyond

Variability in the way organisms reproduce raises numerous, and still unsolved, questions in evolutionary biology. In this study, we emphasize that fungi deserve a much greater emphasis in efforts to address these questions because of their multiple advantages as model eukaryotes. A tremendous diversity of reproductive modes and mating systems can be found in fungi, with many evolutionary transitions among closely related species. In addition, fungi show some peculiarities in their mating systems that have received little attention so far, despite the potential for providing insights into important evolutionary questions. In particular, selfing can occur at the haploid stage in addition to the diploid stage in many fungi, which is generally not possible in animals and plants but has a dramatic influence upon the structure of genetic systems. Fungi also present several advantages that make them tractable models for studies in experimental evolution. Here, we briefly review the unsolved questions and extant hypotheses about the evolution and maintenance of asexual vs. sexual reproduction and of selfing vs. outcrossing, focusing on fungal life cycles. We then propose how fungi can be used to address these long-standing questions and advance our understanding of sexual reproduction and mating systems across all eukaryotes.     

Size‐dependent resource allocation and sex allocation in herbaceous perennial plants

Individuals within a population often differ considerably in size or resource status as a result of environmental variation. In these circumstances natural selection would favour organisms not with a single, genetically determined allocation, but with a genetically determined allocation rule specifying allocation in relation to size or environment. Based on a graphical analysis of a simple evolutionarily stable strategy (ESS) model for herbaceous perennial plants, we aim to determine how cosexual plants within a population should simultaneously adjust their reproductive allocation and sex allocation to their size. We find that if female fitness gain is a linear function of resource investment, then a fixed amount of resources should be allocated to male function, and to post‐breeding survival as well, for individuals above a certain size threshold. The ESS resource allocation to male function, female function, and post‐breeding survival positively correlate if both male and female fitness gains are a saturating function of resource investment. Plants smaller than the size threshold are expected to be either nonreproductive or functionally male only.     

Skewed paternity and sex allocation in hermaphroditic plants and animals

Models predict a reduced allocation to sperm when females preferentially use one of two males' sperm and the males do not know who is favoured. An analogous discounting occurs in plants when their paternity success is skewed by random, non-heritable factors such as location in the population and pollinator behaviour. We present a model that shows that skewed paternity can affect the sex allocation of hermaphrodites, that is it leads to a female-biased investment. The model highlights the close links between local mate competition and sperm competition. We use paternity data from Ficus in order to illustrate that skews in paternity success can lead to a high degree of sibling gamete competition in an apparently open breeding system. Since skews in paternity are ubiquitous in hermaphroditic plants and animals these findings should apply broadly.