The productive and reproductive biology of flowering plants

The life cycles, programme of energy expenditure and allocation to reproduction, and the reproductive efforts of three wildAllium species, i.e.,A. Victorialis ssp.platyphyllum, A. monanthum, andA. Grayi, all native to Japan, were studied and compared. Furthermore, their adaptive strategies were discussed from the point of view of life history strategy. First, the reproductive systems, number of male and female gametes borne, and the number and size of propagules produced were critically investigated. In order to estimate the crude reproductive efficiency (sensu Harper and Ogden, 1970) of these species, sequential harvests were taken and the plants were divided into their component structures, dried and weighed. The quantity of dry weight allocated to sexual or vegetative reproduction was obtained by weighing the seeds, bulbils, or bulblets produced at the end of the season. A. Victorialis ssp.platyphyllum showed a rather low reproductive effort. However, the mean seed output per plant was 34.8±16.8 and the productivity appeared very constant every season. Thus, in the natural populations young plants are borne and recruited every season by means of sexual reproduction. A. monanthum was found to be characterized by annual type dry matter economy. The sexuality and reproductive systems of this species turned out to be extremely complex, and ten different reproductive types were distinguished. The exceedingly low efficiency of sexual reproduction in this species is apparently supplemented by vegetative propagation. The dry matter allocation to daughter bulbs at final harvest was very high; whereas the allocation to sexual reproduction was extremely low. InA. Grayi (a polyploid complex of 4X, 5X, and 6X), a surprisingly high amount of the total annual net assimilate is allocated to the bulbils and bulblets. On the other hand, sexual reproductive effort in this species is exceedingly low, even in obligate amphimictic plants. Thus, the recruitment of individuals in a population of this species appears to be largely dependent on vegetative reproduction. Considering the number of bulbils produced in the scape heads, their dispersibility, germinability, and rapid growth after sprouting, the bulbils evidently possess a function almost comparable to seeds. This species no doubt possesses an adapative strategy to unstable, open habitats exposed to frequent disturbances. It is concluded that the life history strategies of plants, as characterized here in this paper for three wildAllium species, have doubtlessly differentiated by adapting to the respective ecological backgrounds of their habitats.     

The molecular biology of self-incompatibility systems in flowering plants

Self-incompatibility is a common mechanism by which flowering plants can exert some control over the process of fertilization. Typically, the self-incompatibility response involves the recognition and rejection of self-incompatible pollen which leads to a block in self-fertilization and, as a consequence, promotes outcrossing. In recent years, considerable progress has been made in the molecular understanding of several self-incompatibility systems. Interestingly, a common mechanism for self-incompatibility is not employed by all flowering plants, but in fact quite diverse mechanisms have been recruited for the rejection of self-incompatible pollen. In this review, the recent advances in the self-incompatibility systems of the Solanaceae, Papaveraceae, and Brassicaceae will be described as well as some of the molecular work that is emerging for the Poaceae and the heteromorphic self-incompatibility systems.     

The formation and function of the female reproductive tract in flowering plants

In angiosperms, sexual reproduction requires a sperm cell, contained within a pollen tube, to fertilize the egg cell. The pollen tubes are capable of growth but have a difficult journey, as egg cells are buried within the ovary of the carpel. Several tissues, known collectively as the reproductive tract, develop within the carpel to facilitate the journey of the pollen tube. The genes involved in the formation and function of the reproductive tract have largely remained a mystery but are crucial for successful fertilization. This review summarizes recent advances in our understanding of the genetic control of reproductive tract development.     

The occurrence of estrogens in relation to reproductive processes in flowering plants

Summary Estrogens, a group of steroidal sex hormones, were determined in pollen, styles or whole pistils using a highly sensitive and specific radioimmunoassay (RIA) and found to be present at levels between 10^–11 and 10^–10 g/g FW. In the style of Liliun davidii Duch., the levels of total estrogen decreased after self-pollination of the open flower. After bud pollination, which partly overcame incompatibility, the levels of both estradiol and total estrogen increased. The levels of estradiol and total estrogen in the pistils of Brassica pekinensis Rupr. increased after compatible pollination during the first 3 days following pollination and decreased thereafter when the data were expressed on a FW basis. These results suggest that estrogens are to some extent associated with the reproductive processes.     

Review. The strength and genetic basis of reproductive isolating barriers in flowering plants

Speciation is characterized by the evolution of reproductive isolation between two groups of organisms. Understanding the process of speciation requires the quantification of barriers to reproductive isolation, dissection of the genetic mechanisms that contribute to those barriers and determination of the forces driving the evolution of those barriers. Through a comprehensive analysis involving 19 pairs of plant taxa, we assessed the strength and patterns of asymmetry of multiple prezygotic and postzygotic reproductive isolating barriers. We then reviewed contemporary knowledge of the genetic architecture of reproductive isolation and the relative role of chromosomal and genic factors in intrinsic postzygotic isolation. On average, we found that prezygotic isolation is approximately twice as strong as postzygotic isolation, and that postmating barriers are approximately three times more asymmetrical in their action than premating barriers. Barriers involve a variable number of loci, and chromosomal rearrangements may have a limited direct role in reproductive isolation in plants. Future research should aim to understand the relationship between particular genetic loci and the magnitude of their effect on reproductive isolation in nature, the geographical scale at which plant speciation occurs, and the role of different evolutionary forces in the speciation process.     

The reproductive biology of two understory plants in the Atlantic rain forest, Brazil

Outcrossing and sexual reproduction of most flowering plants depends on pollinators. Plant traits likely to be involved in pollinator attraction include flower color, shape, and size. Furthermore, plant or flower density and the temporal flowering pattern may have an effect on reproduction. In this study, we examine the pollination ecology, breeding system, female reproductive output, and germination of two tropical understory species, Stenostephanus lobeliiformis (Acanthaceae) and Besleria melancholica (Gesneriaceae), which differ in these traits. Pollinator observations revealed that the dense flowering S. lobeliiformis with pinkish flowers received a higher diversity of pollinators, but visitor frequency measured as visits per flower per hour was much less (0.1 h^?1) than that to B. melancholica, which has a smaller floral display of dull-colored flowers (1.5 h^?1). Pollination experiments revealed that S. lobeliiformis but not B. melancholica is pollen-limited. In addition, both species are partially self-incompatible and depend on pollinators for outcrossing. Natural fruit set of open-pollinated unmanipulated flowers (control treatment) in both species is 22–26 %. Germination studies indicated inbreeding depression in S. lobeliiformis. We conclude that the pollination ecology of these species is influenced by a broad set of traits and that very different combinations of these traits can be successful in terms of reproduction.     

The origin and function of self-incompatibility in flowering plants

Summary The evolutionary significance of self-incompatibility (SI) traditionally has been linked to reduced inbreeding through enforced outcrossing. This view is founded on the premise that outcrossing reduces inbreeding. It is important, when considering the evolutionary significance of any genetic system, to try to distinguish those factors related to the evolution of, from those related to the maintenance of, the system in question. Three factors are considered important for the maintenance of SI: (1) phylogenetic constraint in species descended from SI ancestors, (2) reduced inbreeding in populations, and (3) fitness benefits to individuals resulting from the avoidance of selfing. I suggest that the first two factors should be rejected when considering the origin of SI (whether one or more origins are hypothesized) and that the increase in individual fitness resulting from the avoidance of self-fertilization among individuals that are heterozygous for deleterious alleles may be sufficient to account for the origin of SI. Self-fertilization in plants (except in species that predominantly self-fertilize) generally results in a reduction in fitness of some individuals due to the increased expression of deleterious or lethal recessive alleles, regardless of the degree of inbreeding in the population or the frequency of the allele in question. Inbreeding is a consequence of population structure in many outcrossing plant species. Complex (multi-locus and multi-allelic) systems of SI exist that reduce inbreeding. However, it is argued that these are derived either from simpler systems of SI that may have very little or no effect on inbreeding, in which case any effect on level of inbreeding is secondary, or are not true self-incompatibility systems and are part of a regulatory system that serves to balance the level of inbreeding and outbreeding. Multi-locus and multi-allelic systems of SI and heteromorphic systems of SI are discussed in terms of derived versus ancestral characteristics. A reassessment of the role of breeding systems in the development of a population structure promoting inbreeding is suggested, which may have been of crucial importance in the success and diversification of angiosperms.     

The reproductive biology of Cyclops vicinus

Investigations of the reproductive biology of Cyclops vicinusrevealed that mated females oscillate between gravid and nongravidreproductive conditions. The gravid condition can be dividedin two recognizable phases: gravid/nonovigerous and gravid/ovigerous,the former phase being shortest. The maturation of new oocytestakes place when the old egg sacs are still being carried; thisensures a rapid clutch succession. Females which remain unmated,extrude few eggs, in no case complete egg sacs, and remain gravidthus conserving oocytes. Females which mate only once, showa similar reproductive pattern (clutch size and clutch succession)to those which remain combined with males, and thus have theopportunity to remate, but tend to produce fewer clutches. Malesare able to fertilize 3–4 females day^–1. Matingcapacity of males is possibly limited by the time needed tofill a new spermatophore. Short-term starvation (5 days) lengthenedclutch-to-clutch periods and diminished clutch size. When thestarvation period started in the gravid/ovigerous phase, a normalclutch was extruded but no new oocytes matured during starvation,indicating that the energy for egg maturations is provided inthe first part of the reproductive cycle.     

The timetable for allopolyploidy in flowering plants

Background

Our understanding of the processes and dynamics of allopolyploid speciation, the long-term consequences of ploidal change, and the genetic and chromosomal changes in new emerged allopolyploids has substantially increased during the past few decades. Yet we remain uncertain about the time since lineage divergence when two taxa are capable of spawning such entities. Indeed, the matter has seemed intractable. Knowledge of the window of opportunity for allopolyploid production is very important because it provides temporal insight into a key evolutionary process, and a temporal reference against which other modes of speciation may be measured.

Scope

This Viewpoint paper reviews and integrates published information on the crossability of herbaceous species and the fertility of their hybrids in relation to species'' divergence times. Despite limitations in methodology and sampling, the estimated times to hybrid sterility are somewhat congruent across disparate lineages. Whereas the waiting time for hybrid sterility is roughly 4–5 million years, the waiting time for cross-incompatibility is roughly 8–10 million years, sometimes considerably more. Strict allopolyploids may be formed in the intervening time window. The progenitors of several allopolyploids diverged between 4 and 6 million years before allopolyploid synthesis, as expected. This is the first study to propose a general temporal framework for strict allopolyploidy. This Viewpoint paper hopefully will stimulate interest in studying the tempo of speciation and the tempo of reproductive isolation in general.     

Self-incompatibility in flowering plants

Self-pollination in some groups of plants is prevented by a sophisticated biochemical signalling system. The molecule active in the female emerges as a highly charged glycoprotein, but the identity of the male determinant remains unknown. Studies of both the molecular biology and the physiology of the interaction suggest that the female polypeptide belongs to a family of glycoproteins which may play an additional, and more general, role in pollination. Pollen compatibility is controlled by one of two genetic systems and new information indicates a mechanism by which they may have arisen, together with the different stigma types with which they are correlated.     

Apomixis and amphimixis in flowering plants

The objective of the present article is to compare apomictic and sexual reproduction (amphimixis) in flowering plants. Light-optical and ultrastructural aspects of the cytoembryological processes in apomicts, beginning with the early stages of development of the ovule and concluding with the newly formed seed, are considered. In the overwhelming majority of apomicts, an inability to develop an autonomous endosperm or to form viable seeds without the involvement of the process of fertilization of the nuclei of the central cell of the embryo sac is observed. Characteristic features of the ultrastructural differentiation of the megasporocytes in diplospory, of aposporous initial cells in apospory, of embryocytes in adventive embryony, and of ovicells in parthenogenesis and synergids in apogamety are identified and are compared to the generative structures of amphimicts. The hypothesis is made that the mechanisms of genetic regulation in the formation and development of the generative structures in apomixis and amphimixis are similar at the cellular level. The present study is not a survey of apomixis in general. Previously published original results that have been obtained by the present author as a result of many years of research in the area of apomixis have served as a basis for the preparation of the study.     

Polyploidy and self-fertilization in flowering plants

Mating systems directly control the transmission of genes across generations, and understanding the diversity and distribution of mating systems is central to understanding the evolution of any group of organisms. This basic idea has been the motivation for many studies that have explored the relationships between plant mating systems and other biological and/or ecological phenomena, including a variety of floral and environmental characteristics, conspecific and pollinator densities, growth form, parity, and genetic architecture. In addition to these examples, a potentially important but poorly understood association is the relationship between plant mating systems and genome duplication, i.e., polyploidy. It is widely held that polyploid plants self-fertilize more than their diploid relatives, yet a formal analysis of this pattern does not exist. Data from 235 species of flowering plants were used to analyze the association between self-fertilization and ploidy. Phylogenetically independent contrasts and cross-species analyses both lend support to the hypothesis that polyploids self-fertilize more than diploids. Because polyploidy and self-fertilization are so common among angiosperms, these results contribute not only to our understanding of the relationship between mating systems and polyploidy in particular, but more generally, to our understanding of the evolution of flowering plants.     

The epigenetic basis of gender in flowering plants and mammals

What makes a sperm male or an egg female, and how can we tell? A gamete's gender could be defined in many ways, such as the sex of the individual or organ that produced it, its cellular morphology, or its behaviour at fertilization. In flowering plants and mammals, however, there is an extra dimension to the gender of a gamete – due to parental imprinting, some of the genes it contributes to the next generation will have different expression patterns depending on whether they were maternally or paternally transmitted. The non-equivalence of gamete genomes, along with natural and experimental modification of imprinting, reveal a level of sexual identity that we describe as ‘epigender’. In this paper, we explore epigender in the life history of plants and animals, and its significance for reproduction and development.