Female frequencies and fitness components between sex phenotypes among gynodioecious populations of the colonizing species Trifolium hirtum All. in California

Summary Male sterility has been recently discovered in Californian populations of rose clover (Trifolium hirtum). This study describes the frequency distribution of male sterility in Turkish and Californian populations, and compares fitness components of hermaphrodites and females. As male-steriles were found in Turkey, it is likely that they were introduced to California during the 1940's with the original material derived from Turkey. The spread of male-sterile genotypes in California has given rise to an asymmetrical frequency distribution of male sterility with positive skewness. The frequency of females has not exceeded fifty percent and it appears to be temporally stable in most of the Californian populations. The hypothesis that female frequencies and fitness differences between phenotypes are correlated was tested by comparing sex phenotypes in seven populations with contrasting levels of male sterility. The analysis of those populations showed no evidence for such a correlation as no significant differences were found between sex phenotypes for fecundity and seed germination. The hypothesis that females are maintained due to fitness differences in the progeny of hermaphrodites and females was experimentally tested in the population with maximum frequency of male-steriles. The results showed no significant differences in the demographic performance of the progenies of hermaphrodites and females. The present results are discussed in terms of the possible mechanism of maintenance of gynodioecy in rose clover.     

Inheritance of male sterility in Trifolium hirtum All

The inheritance of male sterility was studied using a set of intra and interpopulation crosses from Californian populations of rose clover (Trifolium hirtum All.). The F2 gave higher frequencies of male steriles in interpopulation crosses than in intrapopulation crosses. Several F2 families gave ratios that were compatible with models of two and three complementary genes. However, the frequency distribution of male sterility among F3 families rejected such models. Given that single and two gene models were rejected, we interpret the results using a cytoplasmic-nuclear model with more than two restorer loci which is at least consistent with the observed results.     

Recombination and linkage disequilibrium among mitochondrial genes in structured populations of the gynodioecious plant Silene vulgaris

The impact of intergenic recombination on the population genetics of plant mitochondrial genomes is unknown. In an effort to study this in the gynodioecious plant Silene vulgaris three-locus PCR/RFLP genotypes (based on the mitochondrial genes atpA, cox1, and cob) were determined for 239 individuals collected from 20 North American populations. Seventeen three-locus PCR/RFLP genotypes were found. Recombination was indicated by observation of each of the four two-locus genotypes possible when the two most common alleles are considered for each of two loci. Based on these common alleles the absolute values of standardized linkage disequilibrium |D'| between pairs of loci range from 0.17 to 0.78. This indicates modest disequilibrium, rather than the maximum value expected in the absence of recombination |D'=1|, or the linkage equilibrium expected if recombination is pervasive (D'=0). Values of D' did not depend on which pair of loci contributed alleles to the analysis. The direction of D' obtained for the common atpA and cox1 alleles was comparable in sign and magnitude to that obtained by examining similar information obtained in a prior study of European samples. All three loci indicated a high degree of population structure (average FST=0.63), which would limit the within-population genetic diversity required for intergenic recombination to create novel genotypes, if most mating is local. Thus, population structure acts as a constraint on the approach to linkage equilibrium.     

Simple mathematical models to describe the rate of mating in insect populations

Simple models are constructed to describe the rate of mating in insect populations. The models are based on the assumption of random mate-searching in a closed habitat, including four parameters, i.e., population size, sex ratio, searching efficiency and male's capacity on mating frequency. The modes of effects of these parameters on the rate of mating are analyzed and some principles deduced are discussed in relation to the mating process in natural populations.     

Isozyme analysis of genetic variability and population structure of Lactuca L. germplasm

Isozymes were used to investigate the genetic variability, population structure, and relationships of Lactuca germplasm. The isozyme systems revealed 16 putative loci of a total of 31 alleles. Out of these 16 loci, 11 were polymorphic. The average values of expected heterozygosity (H_e), observed heterozygosity (H_o), mean number of alleles per locus (A) and effective number of alleles per locus (A_e) were 0.2227, 0.266, 1.3005 and 1.369, respectively. The average fixation indices were lower than zero for most of the accessions studied, indicating an excess of heterozygotes. Genetic differentiation among accessions (F_ST) exhibited that 51.3% of the isozyme variation was recorded among accessions, and 48.7% of the genetic variation resided within accessions. The average values of total heterozygosity (H_T) and intra-accessional genetic diversity (H_S) were 0.352 and 0.171, respectively. Moreover, the inter-accessional genetic diversity (D_ST) ranged from 0 to 0.424 with an average of 0.18. Cluster analysis revealed that L. sativa cultivars were distributed throughout different Lactuca species. Thereby, isozymes results confirms the hypothesis of the polyphyletic origin of L. sativa. This high level of genetic variation proved that isozymes are efficient for polymorphism analysis of Lactuca germplasm.     

Isozyme variation and differentiation of morphologically cryptic species in the Brachypodium distachyon complex

Isozyme variation of nine enzymes among three morphologically cryptic species of the Brachypodium distachyon complex was studied with PAGE. Aspartate aminotransferase isozymes AAT-A and AAT-C, aryl alcohol dehydrogenase AAD-B, malate dehydrogenase MDH-A and leucine aminopeptidase LAP-A in the allotetraploid B. hybridum display fixed heterozygosities with codominant expression of homoeozymes corresponding to divergent orthozymes of diploids B. distachyon and B. stacei. The multilocus isozyme genotype inherent to most accessions of B. hybridum combines two divergent multilocus isozyme genotypes characteristic of most accessions of B. distachyon and B. stacei. The maternally inherited chloroplast-encoded RUBISCO isozymes show that B. stacei is a maternal progenitor of B. hybridum, whereas the nuclear-encoded isozymes AAD-B, AAT-A, AAT-C, LAP-A, and MDH-A confirm that B. distachyon is the paternal progenitor. These data provide a new isozyme-based support to the karyological and DNA marker data on the allotetraploid nature of B. hybridum and its origin from diploids B. distachyon and B. stacei.     

Population and species variation of minisatellite DNA in Plantago

Three Plantago species were surveyed for within- and between-population variation at DNA sequences detected with the M13 minisatellite probe. The levels and patterns of variation detected by this probe correspond to those expected from the mating systems of the species. The highly-selfing species P. major has a relatively low variability of minisatellite sequences within populations and considerable differentiation between populations. The outcrossing species P. lanceolata exhibits higher minisatellite variability within populations and moderate differentiation between populations. P. coronopus, with a mixed mating system, has levels of variation intermediate between P. major and P. lanceolata. The levels of variation within and between populations corresponds, in general, to the levels of allozyme variation determined in an earlier study. Mating system and population structure appear to have a major influence on M13-detected fragment variation.     

Isozyme variation and species relationships in the genus Lolium L. (ryegrasses,Graminaceae)

Thirty-two natural populations belonging to the eight species of the genus Lolium (ryegrass) or to Festuca pratensis (meadow fescue) were recorded for allelic frequencies at 13 isozyme loci. Cultivated ryegrass (L. perenne and L. multiflorum), meadow fescue, and the annual L. rigidum, are true outbreeders. The other species are true inbreeders, except for L. canariense, which shows a moderate level of cross fertilisation (20%). Hierarchical clustering from Nei's unbiased distance leads to four groups. The three self-pollinating, weed species, L. temulentum, L. remotum and L. persicum, belong to the first cluster, which is the most differentiated one. The second cluster comprises L. multiflorum, L. subulatum and most populations of L. rigidum. All L. perenne populations belong to the third cluster, as do two of L. rigidum. The average genetic distance within the L. perenne group is very low. Surprisingly, the fourth cluster groups together L. canariense and Festuca pratensis. The data suggest that L. rigidum is the species with the greatest diversity, and could be a common ancestor of the genus. Knowledge of historical processes of domestication could help to calibrate the molecular clock.     

Breeding systems in two species of the genus Helleborus (Ranunculaceae)

ABSTRACT

The results of various types of manual and free pollination are reported in Helleborus bocconei and Helleborus foetidus, two sympatric species of the understorey of submediterranean woods that flower in January–March and are pollinated by bumble-bees of the genus Bombus. Both species were found to be partially self-compatible even if at different rates; geitonogamy seems to be more frequent in H. foetidus and spontaneous self-pollination does not usually occur for morphological reasons, and the species are not apomictic. Efficiency in seed production after free pollination was very high indicating that in spite of harsh climatic condition, there were no limitations imposed by pollen or pollinators.     

Population characteristics of endangered species Taxus chinensis var. mairei and its conservation strategy in Shanxi, China

The conservation of the endemic, endangered, and first-class protected tree species, Taxus chinensis var. mairei (Lem. & H. Lev.) W. C. Cheng & L. K. Fu, is urgent in China. To assist the protection of this important medicinal plant species, the biological and ecological characteristics of its populations were investigated and analyzed in Shanxi. The study area, southeastern Shanxi, is a special region suitable for this species growth and development. The floristic composition, life form spectrum, and species diversity of T. chinensis var. mairei forests reflected the features of both warm-temperate and subtropical regions. Size classes in DBH and static life table analysis showed that the population structure of T. chinensis var. mairei was of the increase-type. Plant density, survivorship, and mortality decreased with increase in age. Distribution patterns of individuals were basically clumped and saplings tended to be distributed randomly. Disturbance from collection of leaves, branches, and stems for medicine was an important factor affecting population structure, dynamics, and distribution. The sex ratio of female/male and the efficiency of sexual reproduction were very low. The main way of reproduction for this species is vegetative propagation. Based on these characteristics, some suggestions, such as establishing nature reserves, improving population structure by cultivation and transplantation, developing seed gardens, and plantations, etc., for the conservation management of T. chinensis var. mairei and its forests in Shanxi are discussed.     

The mating system in natural and shelterwood stands of Douglas-fir

Summary Mating systems in two pairs of old-growth uncut and adjacent shelterwood stands of Douglas-fir (Pseudotsuga menziesii var. menziesii) were compared by estimating the proportions of viable progenies due to outcrossing (t) with both single-locus and multilocus techniques. Single-locus population estimates (_s), ranging from 0.41 to 1.16, were significantly (P<0.05) heterogeneous among loci in three of four stands; mean single-locus estimates for shelterwoods were not significantly different from those for uncut stands. Multilocus population estimates (_m) ranged from 0.94 to 1.00; again, estimates for shelterwoods were not significantly different from those for uncut stands. Multilocus estimates were slightly higher than mean single-locus estimates for uncut stands but were nearly equivalent for shelterwoods, suggesting that related matings other than selfs may be associated with uncut stands, but not shelterwoods. Individual-tree outcrossing rates (_{m i}), estimated for six trees in each shelterwood, ranged from 0.90 to 1.10 and were significantly heterogeneous among trees at one of the two shelterwoods. Outcrossing was high (> 0.90) in both uncult and shelterwood stands, and no evidence indicated that low parent-tree density had affected stand outcrossing rates.FRL 1918, Forest Research Laboratory. Oregon State University, Corvallis, USA     

Breeding systems and population structure in Limnanthes

Summary The breeding systems of seven Limanthes (Limanthaceae) populations, including one inbreeding and three outbreeding taxa, were quantified using a multilocus outcrossing rate estimator (t_m) and autofertility estimates. Along with the assays of heterozygosity levels, these data were used to separate components of effective outcrossing in terms of Wright's equilibrium inbreeding coefficient (F_e) and adult (F_A) and zygotic (F_Z) fixation indices. The patchy distribution of alleles as a potential source of substructure inbreeding was tested from the allelic frequencies mapped along a linear transect. Evidence for consanguineous matings in restricted neighborhoods and for selection at two different life cycle stages, and the efficiency of the protandrous breeding system were noted and discussed. Multilocus estimates of outcrossing are useful for their greater precision and unbiased nature while single locus estimates can help in detecting the effects of selection and population substructure. The data generally support the heterozygosity paradox noted by Brown (1979) but further suggest that the paradox may often result from a lack of precision of outcrossing estimates and from overlooking the stages of the life cycle being sampled.     

Parasitic systems and the structure of parasite populations

 The analysis of population systems is carried out on the basis of the spatial and functional classification of populations developed by V.N. Beklemishev. The population system is a functional part of a particular community. Steady interrelationships between population systems of different species within the community (referred to as ”community links”) appear to be a prerequisite for the formation of a complex of population systems. A prominent example of this is the parasitic system. The parasitic system is the population system of a parasite with all the connected populations of its hosts. The complexity of a parasitic system depends on: (1) peculiarities of the life cycle of the parasite, since its population system is the organizing component of the parasitic system and (2) subdivision of the environment for the parasites. The first trait is discussed from the standpoint of the phase structure of populations, which is clearly seen in parasites. The second one comprises the organization of the parasites’ environment according to the scale of variability (interspecies, interpopulation or intrapopulation) of hosts. These make it possible to recognize spatial and functional parts in the framework of the parasitic system. A critical review of the terminology is presented together with a list of the pertaining vocabulary. Received: 15 January 1998 / Accepted: 15 October 1998     

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.     

Genetic structure of black abalone (Haliotis cracherodii) populations in the California islands and central California coast: Impacts of larval dispersal and decimation from withering syndrome

The genetic structure of black abalone populations in the southern California Islands and central California coast was investigated by protein electrophoresis. Detailed sampling of San Nicolas Island (SN) permitted investigation of microgeographic genetic differentiation among local geomorphological features. In addition, temporal genetic differentiation was assessed by comparing juveniles and adults at three islands, San Miguel (SM), Santa Cruz (SC) and San Nicolas (SN). Mainland and island locations were genetically differentiated based on allele frequency differences and the presence of private alleles in some island populations. Although microgeographic genetic structure among sites on SN was weak and not statistically significant, heterozygosity varied among sites, with diversity decreasing from west to east. In addition, investigation among cohorts showed that adults were genetically differentiated among island locations, whereas no differences among juveniles were detected. Genetic differentiation between adult and juvenile abalones was detected at SC but not SM or SN. These data are generally consistent with local recruitment augmented by relatively more gene flow among island populations than among island and mainland populations, and possible selection acting on immigrant recruits.     

Pollen limitation and distance-dependent fecundity in females of the clonal gynodioecious herb Glechoma hederacea (Lamiaceae)

Summary Pollen movement is often restricted in natural populations, and insufficient pollination is a potential constraint on sexual reproduction in outcrossing species. Seed-set should decrease with increased distance from the pollen source in outcrossing plants. This prediction was tested using females of the clonal, gynodioecious herb Glechoma hederacea in three natural populations. In controlled pollinations, both hermaphrodites and females had similar high percentages of fruit-set and seed-set. In a natural population where a female clone was isolated from the nearest hermaphroditic clone by c. 100 m, fruit-set was low (1%). In another population where hemaphroditic clones were rare and female clones had a patchy distribution, fruit-and seed-set in females were pollen-limited and decreased with increased distance from the nearest pollen source. The estimated mean pollen dispersal distance was 5.9 m when calculated on fruit-set and 5.3 m when calculated on seed-set. The most frequent pollinators were bumblebees. The mean and median distances moved by pollinators between ramets were 0.13 m and 0.05 m. In a third population where female clones were isolated from the nearest hermaphrodites by more than 200 m, fruit-set was 0%. After introduction of 16 hermaphroditic ramets in the center of the female clone, fruit-set varied between 0% and 100% in individual female ramets. Fruit-set decreased with increased distance from the pollen source. The mean and median pollen movement distances were 1.06 m and 0.54 m.     

The evolution of self-compatible and self-incompatible populations in a hermaphroditic perennial, <Emphasis Type="Italic">Trillium camschatcense</Emphasis> (Melanthiaceae)

The evolution of selfing from outcrossing ancestors is known to have occurred repeatedly in angiosperms. Theoretical studies have argued that the transition from outcrossing to selfing is accomplished more easily than the reverse case, and phylogenetic analyses involving self-compatible (SC) and self-incompatible (SI) species has basically supported this assumption. The evolutionary direction of self-compatibility and self-incompatibility was examined in Trillium camschatcense, which contains geographically widespread SC populations, and restricted SI populations. Ecological surveys have revealed that the SC populations were suitable for outcrossing, and selfing in these populations did not confer any fitness advantage. Since reproductive fitness indicates the possibility of an evolutionary shift from self-compatibility to self-incompatibility, the phylogenetic relationships of SI and SC populations of T. camschatcense were investigated based on cpDNA variations and nuclear DNA microsatellite polymorphisms. Although phylogenetic analyses did not provide credible evidence to determine evolutionary direction, the SI populations turned out to be monophyletic with extremely low genetic differentiation. Based on these results, we proposed two possible scenarios for the evolutionary backgrounds of SI and SC populations in T. camschatcense. The plausibility of each scenario was evaluated based on the reproductive and geographical features of the mating systems. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Evolution of male sterility mechanisms in gynodioecious and dioecious species ofCirsium (Cynareae,Compositae)

The genusCirsium comprises both gynodioecious and dioecious species. The observation of microsporogenesis in female plants ofC. montanum, C. oleraceum, C. palustre andC. spinosissimum shows that the male sterility is due to a degeneration of the tapetum. This degeneration occurs more or less early according to the species and, in the light of these results, a scheme of evolution in the male sterility mechanism is proposed. Furthermore, the male sterility mechanism inC. montanum is very similar to that previously found in female plants of the dioecious speciesC. arvense. This fact enhances the possibility of evolution of the dioecy ofC. arvense from the gynodioecy found in other species. According to these results, a general scheme of evolution of sexes in the genusCirsium is proposed.     

Low genetic diversity and significant structuring in the endangered Mentha cervina populations and its implications for conservation

Eighteen populations of the endangered aromatic and medicinal plant Mentha cervina (Lamiaceae) were sampled across its natural range, in the western half of the Iberian Peninsula, and inter-simple sequence repeats (ISSRs) markers were used to assess genetic diversity and population structure. M. cervina populations exhibited a relatively low genetic diversity (percentage of polymorphic loci PPB = 14.2–58.3%, Nei's genetic diversity H_e = 0.135–0.205, Shannon's information index I = 0.08 − 0.33). However, the genetic diversity at species level was relatively high (PPB = 98.3%; H_e = 0.325; I = 0.23). The results of the analysis of molecular variance indicated very structured populations, with 50% of the variance within populations, 44% among populations and 6% between regions defined by hydrographic basins, in line with the gene differentiation coefficient (G_ST = 0.532). A Mantel test did not find significant correlation between genetic and geographic distance matrices (r = 0.064), indicating that isolation by distance is not shaping the present genetic structure. The levels and patterns of genetic diversity in M. cervina populations were assumed to result largely from a combination of evolutionary history and its unique biological traits, such as breeding system, low capacity of dispersion, small effective size and habitat fragmentation. The high genetic differentiation among populations indicates the necessity of conserving the maximum possible number of populations. The results also provide information to select sites for ex situ conservation. Optimal harvesting strategies, cultivation and tissue culture should also be developed as soon as possible to guarantee sustainable use of the species under study.