Floral sex ratio variation in hermaphrodites of gynodioeciousChionographis japonica var.kurohimensis Ajima et Satomi (Liliaceae)

Intrapopulation and interpopulation variations in floral sex ratio in hermaphrodites of gynodioeciousChionographis japonica var.kurohimensis (Liliaceae) were examined. The relative ratio of male flowers to total flowers (male and perfect flowers) decreased with plant size, suggesting size-dependent gender modification. The relative ratio of male flowers per population-basis is negatively correlated with the mean number of perfect flowers. Since the number of perfect flowers proportionally increased with plant size, populations showing low maleness consist of relatively bigger plants and are considered to be in high-quality environment. On the other hand, the relative ratio of male flowers per population basis is independent of female frequency in the population. Plasticity in gender expression probably plays an important role of maintenance of gynodioecy inC. japonica var.kurohimensis.     

Nuclear androdioecy and gynodioecy

We formulate two single-locus Mendelian models, one for androdioecy and the other one for gynodioecy, each with 3 parameters: t the male (female) fertility rate of males (females) to hermaphrodites, s the fraction of the progeny derived from selfing; and g the fitness of inbreeders. Each model is expressed as a transformation of a 3 dimensional zygotic algebra, which we interpret as a rational map of the projective plane. We then study the dynamics for the evolution of each reproductive system; and compare our results with similar published models. In this process, we introduce a general concept of fitness and list some of its properties, obtaining a relative measure of population growth, computable as an eigenvalue of a mixed mating transformation for a population in equilibrium. Our results concur with previous models of the evolution of androdioecy and gynodioecy regarding the threshold values above which the sexual polymophism is stable, although the previous models assume constant the fraction of ovules from hermaphrodites that are self pollinated, while we assume constant the fraction of the progeny derived from selfing. A stable androdioecy requires more stringent conditions than a stable gynodioecy if the amount of pollen used for selfing is negligible in comparison with the total amount of pollen produced by hermaphrodites. Otherwise, both models are identical. We show explicitly that the genotype fitnesses depend linearly on their frequencies. Simulations show that any population not at equilibrium always converges to the equilibrium point of higher fitness. However, at intermediate steps, the fitness function occasionally decreases.     

Inbreeding avoidance and the evolution of gender dimorphism in Wurmbea biglandulosa (Colchicaceae)

How females establish in populations of cosexuals is central to understanding the evolution of gender dimorphism in angiosperms. Inbreeding avoidance hypotheses propose that females can establish and be maintained if cosexual fitness is reduced because they self-fertilize, and their progeny express inbreeding depression. Here we assess the role of inbreeding avoidance in maintaining sexual system variation in Wurmbea biglandulosa. We estimated costs of self-pollination, mating patterns, and inbreeding depression in gender monomorphic (cosexuals only) and dimorphic (males and females) populations. Costs of selfing, estimated from seed set of experimentally self- and cross-pollinated flowers, were severe in both males and cosexuals (inbreeding depression, sigma = 0.86). In a field experiment, intact males that could self produced fewer seeds than both emasculated males and females, whereas seed set of intact and emasculated cosexuals did not differ. Thus, pollinator-mediated selfing reduces fitness of males but not cosexuals under natural conditions. Outcrossing rates of males revealed substantial selfing (t = 0.68), whereas females and cosexuals were outcrossed (0.92 and 0.97). For males, progeny inbreeding coefficients exceeded parental coefficients (0.220 vs. 0.009), whereas for females and cosexuals these coefficients did not differ and approached zero. Differences in coefficients between males and their progeny indicate that selfed progeny express severe inbreeding depression (sigma = 0.93). Combined with inbreeding depression for seed set, cumulative sigma = 0.99, indicating that most or all selfed zygotes fail to reach reproductive maturity. We propose that present sexual system variation in W. biglandulosa is maintained by high inbreeding depression coupled with differences in selfing rates among monomorphic and dimorphic populations.     

Maintenance of gynodioecy in Wurmbea biglandulosa (Colchicaceae): gender differences in seed production and progeny success

 In gynodioecious species, females contribute genes to future generations only through ovules, and to persist in populations they must have a compensatory advantage compared with hermaphrodites that reproduce via ovules and pollen. This compensation can result from greater fecundity and/or superior success of progeny from females. We examined differences in seed production and progeny success between females and hermaphrodites in the geophyte Wurmbea biglandulosa to explain the maintenance of females. Females produced more ovuliferous flowers and had more ovules per flower than did hermaphrodites but this did not necessarily result in greater fecundity, in part because seed production of females was pollen-limited. Over four years in one population, open-pollinated females produced 1.32 more seeds than open-pollinated hermaphrodites (range 1.09–1.63). In two other populations examined for one year only females produced 1.07 and 0.79 as many seeds as hermaphrodites. Seed production of open-pollinated females and hermaphrodites was only 55% and 73% that of cross-pollinated plants, respectively, indicating that both genders were pollen-limited but females more so than hermaphrodites. Open-pollinated seeds from females were 1.18–1.27 times more likely to germinate than seeds from hermaphrodites. No gender differences existed in seedling growth or survival. Hermaphrodites were self-compatible, but selfed seed set was only 80% that of crossed seed set. Crossed seed set of females and hermaphrodites did not differ. Assuming nuclear control of male sterility, relative female fitness is insufficient to maintain females at their current frequencies of 17%, and substantial female fitness advantages at later life-cycle stages are required. Received May 4, 2001 Accepted February 25, 2002     

Mitochondrial Genome Dynamics in Plants and Animals: Convergent Gene Fusions of a MutS Homologue

Mitochondrial processes influence a broad spectrum of physiological and developmental events in higher eukaryotes, and their aberrant function can lead to several familiar disease phenotypes in mammals. In plants, mitochondrial genes directly influence pollen development and the occurrence of male sterility in natural plant populations. Likewise, in animal systems evidence accumulates to suggest important mitochondrial functions in spermatogenesis and reproduction. Here we present evidence for a convergent gene fusion involving a MutS-homologous gene functioning within the mitochondrion and designated Msh1. In only plants and soft corals, the MutS homologue has fused with a homing endonuclease sequence at the carboxy terminus of the protein. However, the endonuclease domains in the plants and the soft corals are members of different groups. In plants, Msh1 can influence mitochondrial genome organization and male sterility expression. Based on parallels in Msh1 gene structure shared by plants and corals, and their similarities in reproductive behavior, we postulate that this convergent gene fusion might have occurred in response to coincident adaptive pressures on reproduction. [Reviewing Editor: Dr. Deborah Charlesworth]     

Allozyme variation in Japanese species ofChionographis (Liliaceae)

Allozyme variation was examined in three diploid taxaChionographis japonica var.japonica, var.kurokamiana, andC. koidzumiana and three tetraploid taxaC. japonica var.kurohimensis, ssp.hisauchiana, and ssp.minoensis. Results show thatC. japonica var.kurokamiana is genetically closer toC. koidzumiana than to var.japonica. In the tetraploid taxa, fixed heterozygosities were found at several loci, and this supports the hypothesis that these taxa are allotetraploids. Furthermore, the tetraploid taxa have many unique alleles not found in the diploid taxa. This suggests that sufficient time has passed since the origin of tetraploids for new mutations to have been fixed.     

Male‐biased hermaphrodites in a gynodioecious shrub,Daphne jezoensis

Gynodioecy, a state where female and hermaphrodite plants coexist in populations, has been widely proposed an intermediate stage in the evolutionary pathway from hermaphroditism to dioecy. In the gynodioecy–dioecy pathway, hermaphrodites may gain most of their fitness through male function once females invade populations. To test this prediction, comprehensive studies on sex ratio variation across populations and reproductive characteristics of hermaphrodite and female phenotypes are necessary. This study examined the variation in sex ratio, sex expression, flower and fruit production and sexual dimorphism of morphological traits in a gynodioecious shrub, Daphne jezoensis, over multiple populations and years. Population sex ratio (hermaphrodite:female) was close to 1:1 or slightly hermaphrodite‐biased. Sex type of individual plants was largely fixed, but 15% of plants changed their sex during a 6‐year census. Hermaphrodite plants produced larger flowers and invested 2.5 times more resources in flower production than female plants, but they exhibited remarkably low fruit set (proportion of flowers setting fruits). Female plants produced six times more fruits than hermaphrodite plants. Low fruiting ability of hermaphrodite plants was retained even when hand‐pollination was performed. Fruit production of female plants was restricted by pollen limitation under natural conditions, irrespective of high potential fecundity, and this minimised the difference in resources allocated to reproduction between the sexes. Negative effects of previous flower and fruit production on current reproduction were not apparent in both sexes. This study suggests that gynodioecy in this species is functionally close to a dioecious mating system: smaller flower production with larger fruiting ability in female plants, and larger flower production with little fruiting ability in hermaphrodite plants.     

Parental effects and gender specialization in a tropical heterostylous shrub

Abstract.— Male sterility in hermaphroditic species may represent the first step in the evolution toward dioecy. However, gender specialization will not proceed unless the male-sterile individuals compensate for fitness lost through the male function with an increase in fitness through the female function. In the distylous shrub Erythroxylum havanense , thrum plants are partially male-sterile. Using data collected throughout eight years, we investigated whether thrum individuals have an increased performance as female parents, thereby compensating for their loss of male fitness. We found that thrum plants outperformed pins in the probabilities of seed maturation and germination and long-term growth of the seedlings. In turn, pollen from pin plants achieved greater pollen tube growth rates. Our results suggest that the superior performance of the progeny of thrum maternal plants is a consequence of better seed provisioning via effects of the maternal environment, cytotype or nuclear genes. Overall, our results suggest that E. havanense is evolving toward a dioecious state through a gynodioecious intermediate stage. This evolutionary pathway is characterized by an unusual pattern of gender dimorphism with thrums becoming females and pins becoming males. We propose that this pattern may be better explained by the interaction between male-sterility cytoplasmic genes and the heterostyly supergene.     

Exploring the genetic basis and proximate causes of female fertility advantage in gynodioecious Thymus vulgaris

In many gynodioecous species, females produce more viable seeds than hermaphrodites. Knowledge of the relative contribution of inbreeding depression in hermaphrodites and maternal sex effects to the female fertility advantage and the genetic basis of variation in female fertility advantage is central to our understanding of the evolution of gender specialization. In this study we examine the relative contribution of inbreeding and maternal sex to the female fertility advantage in gynodioecious Thymus vulgaris and quantify whether there is genetically based variation in female fertility advantage for plants from four populations. Following controlled self and outcross (sib, within-population, and between-population) pollination, females had a more than twofold fertility advantage (based on the number of germinating seeds per fruit), regardless of the population of origin and the type of pollination. Inbreeding depression on viable seed production by hermaphrodites occurred in two populations, where inbreeding had been previously detected. Biparental inbreeding depression on viable seed production occurred in three of four populations for females, but in only one population for hermaphrodites. Whereas the maternal sex effect may consistently enhance female fertility advantage, inbreeding effects may be limited to particular population contexts where inbreeding may occur. A significant family x maternal sex interaction effect on viable seed production was observed, illustrating that the extent of female fertility advantage varies significantly among families. This result is due to greater variation in hermaphrodite (relative to female) seed fertility between families. Despite this genetic variation in female fertility advantage and the highly female biased sex ratios in populations of T. vulgaris, gynodioecy is a stable polymorphism, suggesting that strong genetic and/or ecological constraints influence the stability of this polymorphism.     

Identification and mapping of RAPD and RFLP markers linked to a fertility restorer gene for a new source of cytoplasmic male sterility in Beta vulgaris ssp. maritima

 The present study shows that the recently described mitochondrial H haplotype is associated with cytoplasmic male-sterility (CMS). This new source of CMS appears to be different from the mitotype E-associated CMS most frequently found in natural populations. A mitotype H progeny with a sexual phenotype segregation was used to identify a gene restoring male fertility (R1H ). Using bulk segregant analysis (BSA), nine RAPD markers linked to this restorer locus were detected and mapped. The comparison with other Beta genetic maps shows that the closest RAPD marker, distant from R1H by 5.2 cM, belongs to the same linkage group as the monogermy locus. In order to determine the position of R1H more precisely, four RFLP loci within this linkage group were mapped in the segregating progeny. It thus became possible to construct a linkage map of the region containing the RFLP, RAPD and R1H loci. The closest RFLP marker was located 1.7 cM away from R1H. However, a nuclear gene restoring the ‘Owen’ CMS which is currently used in sugar beet breeding is reportedly linked to the monogermy locus, raising the question of a possible identity between the new CMS system and the ‘Owen’ CMS. Received: 15 September 1997 / Accepted: 1 December 1997