The control of incompatibility in distylous Pulmonaria affinis Jordan (Boraginaceae)

In P. affinis, pin pollen is shorter on average than thrum pollen. Pins have more coronal hooks on stigmatic papillae than thrums, but gaps between papillae are relatively smaller for thrums. Pin stigmas receive more pollen than thrum stigmas. Thrum stigmas receive more (dissortive) pin pollen, but pin stigmas are assortively pollinated. Pollen only germinates when trapped below papilla coronas. On thrum stigmas, most trapped pollen is pin. Pollen germination is better on thrum stigmas than pin stigmas, and thrum stigmas show a close relationship between numbers of legitimate pollen grains, numbers of germinating grains, and numbers of pollen tubes in the style. There is no inhibition of illegitimate pollen germination. Illegitimate pollen tubes are inhibited in the style. Incompatibility operates by a combination of dissortive pollination, dissortive pollen trapping, and stylar pollen tube inhibition. All heteromorphic features differing between pins and thrums are implicated in the inhibition of within-morph fertilization in thrums.     

The effect of color polymorphism on mortality in the aphidMacrosiphoniella yomogicola

We examined body color polymorphism in the aphidMacrosiphoniella yomogicola from July to September 1993. We classified body color into eight types: green 1, green 2, red 1, red 2, white, orange, yellow and mist. The frequencies of body color varied with time and among patches of the host plant, yomogi (Artemisia spp.). Color diversity within a shoot was calculated using the Shannon diversity index. Of five usable data sets, three showed negative relationships between color diversity and mortality. The regression coefficients for two of these relationships were significant. No significant relationship between mortality and the number of aphids was found. The color diversity was not significantly related to a particular body color found on a yomogi shoot. Color polymorphism may be maintained because selection may favor a high color diversity on the host plant shoot.     

Adaptive evolution of sexual systems in pedunculate barnacles

How and why diverse sexual systems evolve are fascinating evolutionary questions, but few empirical studies have dealt with these questions in animals. Pedunculate (gooseneck) barnacles show such diversity, including simultaneous hermaphroditism, coexistence of dwarf males and hermaphrodites (androdioecy), and coexistence of dwarf males and females (dioecy). Here, we report the first phylogenetically controlled test of the hypothesis that the ultimate cause of the diverse sexual systems and presence of dwarf males in this group is limited mating opportunities for non-dwarf individuals, owing to mating in small groups. Within the pedunculate barnacle phylogeny, dwarf males and females have evolved repeatedly. Females are more likely to evolve in androdioecious than hermaphroditic populations, suggesting that evolution of dwarf males has preceded that of females in pedunculates. Both dwarf males and females are associated with a higher proportion of solitary individuals in the population, corroborating the hypothesis that limited mating opportunities have favoured evolution of these diverse sexual systems, which have puzzled biologists since Darwin.     

The evolution of repeated mating under sexual conflict

In insects, repeated mating by females may have direct effects on female fecundity, fertility, and longevity. In addition, a female's remating rate affects her fitness through mortality costs of male harassment and ecological risks of mating such as predation. We analyse a model where these female fitness factors are put into their life-history context, and traded against each other, while accounting for limitations because of mate availability. We solve analytically for the condition when female multiple mating will evolve. We show that the probability that a female mates with a courting male decreases with increases in population density. The extent of conflict between the sexes thus automatically becomes larger at higher densities. However, because at higher densities females meet males at a higher rate, the resulting ESS female remating rate is independent of population density. The female remating probability is in conflict with male adaptations that increase male mating rate by persuading or forcing females to mate, and also in conflict with male adaptations for protecting the own sperm from being removed by future female mates. We show that the relative importance of these conflicts depends on population density.     

Female reproductive success and the evolution of mating-type frequencies in tristylous populations

In tristylous populations, mating-type frequencies are governed by negative frequency-dependent selection typically resulting in equal morph ratios at equilibrium. However, Narcissus triandrus generally exhibits long-styled (L)-biased populations with a deficiency of the mid-styled (M)-morph. Here we used a pollen-transfer model and measurements of female fertility in natural populations to investigate whether these uneven morph ratios were associated with variation in female reproductive success. Our theoretical analysis demonstrated that morph ratio bias can result from maternal fitness differences among the morphs, and that these effects were magnified by asymmetrical mating. In nine out of 15 populations of N. triandrus, seed set differed significantly among the morphs, but pollen limitation occurred in only two of 11 populations investigated. Average seed set of the M-morph was positively associated with its frequency in populations. Flower size was negatively correlated with the seed set of the M-morph. Our results suggest that interactions between mating patterns and female fertility are responsible for variation in morph frequencies and loss of the M-morph from tristylous populations of N. triandrus.     

New perspectives on the evolution of plant mating systems

BACKGROUND: The remarkable diversity of mating patterns and sexual systems in flowering plants has fascinated evolutionary biologists for more than a century. Enduring questions about this topic include why sexual polymorphisms have evolved independently in over 100 plant families, and why proportions of self- and cross-fertilization often vary dramatically within and among populations. Important new insights concerning the evolutionary dynamics of plant mating systems have built upon a strong foundation of theoretical models and innovative field and laboratory experiments. However, as the pace of advancement in this field has accelerated, it has become increasingly difficult for researchers to follow developments outside their primary area of research expertise. SCOPE: In this Viewpoint paper we highlight three important themes that span and integrate different subdisciplines: the changes in morphology, phenology, and physiology that accompany the transition to selfing; the evolutionary consequences of pollen pool diversity in flowering plants; and the evolutionary dynamics of sexual polymorphisms. We also highlight recent developments in molecular techniques that will facilitate more efficient and cost-effective study of mating patterns in large natural populations, research on the dynamics of pollen transport, and investigations on the genetic basis of sexual polymorphisms. This Viewpoint also serves as the introduction to a Special Issue on the Evolution of Plant Mating Systems. The 15 papers in this special issue provide inspiring examples of recent discoveries, and glimpses of exciting developments yet to come.     

Inbreeding depression in two species of Mimulus (Scrophulariaceae) with contrasting mating systems

We examined the effect of self- and cross-pollination on germination success, flowering probability, pollen and ovule production, survivorship, and adult aboveground biomass in two species of Mimulus with contrasting mating systems: the highly seifing M. micranthus and an outcrossing population of M. guttatus. Cross-pollinations were performed both within and between populations in order to examine the scale at which the genetic load is distributed. We found significant inbreeding depression in M. guttatus in four of the six traits, with the highest inbreeding depression observed in biomass (68% and 69% based on within- and between-population crosses, respectively) and lowest in ovule production (21% based on between-population crosses only). M. micranthus displayed significant inbreeding depression in only two of the six traits examined. Again, we observed the highest inbreeding depression in biomass (47–60% based on within- and between-population crosses, respectively), but both traits showing significant differences between self and outcross progeny expressed lower inbreeding depression than in M. guttatus. We detected no significant inbreeding depression for either pollen or ovule production in M. micranthus. An estimate of total inbreeding depression based on the multiplicative effects of all traits was also lower in M. micanthus than∗∗∗ in M. guttatus. Our results are consistent with the expected purging of genetic load in populations with high selfing rates. The absence of inbreeding depression in M. micranthus pollen and ovule production, two traits with strong links to fitness in a selfing annual, further suggests the important role of directional selection in determining the population's genetic load. Comparison of cross-pollinations made within and between populations revealed little evidence of divergence of genetic load among the M. micranthus and M. guttatus populations examined.     

Clonal growth is enhanced in the absence of a mating morph: a comparative study of fertile stylar polymorphic and sterile monomorphic populations of Nymphoides montana (Menyanthaceae)

Background and Aims

Many aquatic species with stylar polymorphisms have the capacity for clonal and sexual reproduction and are sensitive to the balance of the two reproductive modes when there are a limited number of mating morphs within a population. This study asked how the clonal and sexual reproductive modes perform in populations that contain only a single morph and where fitness gain through sexual reproduction is rare. In clonal aquatic Nymphoides montana, polymorphic populations normally contain two mating morphs in equal frequencies. Populations are sexually fertile and appear to be maintained by pollen transfer between the two partners. However, in a monomorphic population of N. montana where mating opportunities are unavailable, female and male function is impaired and clonality maintains the population. Here, the consequences of intraspecific variation in sexuality were explored between monomorphic and polymorphic N. montana populations in eastern Australia.

Methods

Comparative measurements of male and female fertility, total dry mass and genotypic diversity using ISSR markers were made between populations with variable sexuality.

Key Results and Conclusions

Very few seeds were produced in the monomorphic population under natural and glasshouse conditions due to dysfunctional pollen and ovules. Stigma–anther separation was minimal in the monomorphic population, which may be a consequence of the relaxed selective pressures that regulate the maintenance of sexual function. However, clonal reproduction was favoured at the expense of sexual reproduction in the monomorphic population; this may facilitate the establishment of sterility throughout the population via resource reallocation or pleiotropic effects. The ISSR results showed that the monomorphic population was one large, single genotype, unlike the multi-genotypic fertile polymorphic populations. Evolutionary loss of sex in a clonal population in which a mating morph is absent was evident; under these conditions clonal growth may assure reproduction and expand the population via spreading stolons.     

Frequency-dependent selection by pollinators: mechanisms and consequences with regard to behaviour of bumblebees Bombus terrestris (L.) (Hymenoptera: Apidae)

Bombus terrestris , a typical pollinating insect species, was offered artificial flowers of two different corolla colours with the same sucrose solution reward in an array. Common colours were significantly preferred, and the strength of the frequency-dependent response increased as a result of learning. There were also frequency-independent biases towards blue flowers, probably because blue flowers appeared more conspicuous to bumblebees than yellow flowers, and the degree of preference for blue was greater when flowers had low nectar rewards. Flower-to-flower movements by individual bumblebees between flowers were non-random, were biased to movements within the same flower colour, and were also dependent on morph frequency. The mechanisms governing flower selection in bumblebees are discussed. Pollinators foraging similarly in a natural situation would induce positive frequency-dependent selection, assortative mating, and directional selection on different corolla colour morphs of the plant population being visited, resulting in stabilizing selection for a single flower colour.     

Pollen competition reduces inbreeding depression in Collinsia heterophylla (Plantaginaceae)

We tested two predictions of the hypothesis that competition between self-pollen may mitigate negative genetic effects of inbreeding in plants: (1) intense competition among self-pollen increases offspring fitness; and (2) pollen competition reduces the measured strength of inbreeding depression. We used Collinsia heterophylla (Plantaginaceae), an annual with a mixed mating system, to perform controlled crosses in which we varied both the size of the pollen load and the source of pollen (self vs. outcross). Fitness of selfed offspring was higher in the high pollen-load treatment. Our second prediction was also upheld: inbreeding depression was, on average, lower when large pollen loads were applied (11%) relative to the low pollen-load treatment (28%). The reduction was significant for two fitness components relatively late in the life-cycle: number of surviving seedlings and pollen-tube growth rate in vitro. These findings suggest that intermittent inbreeding, which leads to self-fertilization in plants with genetic loads, may select for traits that enhance pollen competition.     

Reproductive correlates of mating system variation in Eichhornia paniculata (Spreng.) Solms (Pontederiaceae)

Comparative studies of related plant species indicate that evolutionary shifts in mating systems are accompanied by changes in reproductive attributes such as flower size, floral morphology, and pollen/ovule ratio. Recent theoretical work suggests that patterns of investment in reproduction should also change with the mating system. In a glasshouse study, we investigated the extent to which mating system differences among populations of Eichhornia paniculata (Pontederiaceae) were correlated with changes in allocation to male and female function, floral display, and the regulation of investment in reproduction through fruit and ovule abortion. Significant differences in the amount of biomass allocated to reproductive structures were evident among six populations of E. paniculata. As predicted by sex allocation theory, the proportion of dry weight allocated to male function decreased with the outcrossing rate of populations. Six of the eight attributes used to characterize floral display also differed significantly among populations. However, with the exception of two attributes describing the number of flowers produced by inflorescences, these were not correlated with outcrossing rate. Levels of fruit and ovule abortion were determined in two populations with contrasting mating systems under different nutrient and pollination treatments. Virtually all fruits initiated by plants from a self-fertilizing population were matured, while the amount of fruit abortion in an outcrossing population increased with flower production. Ovule abortion was low in both populations. Our results demonstrate that the evolution of self-fertilization in E. paniculata is associated with changes in investment to reproduction that normally distinguish selfing and outcrossing species.     

The mismeasurement of sexual selection

Sexual selection can explain major micro‐ and macro‐evolutionary patterns. Much of current theory predicts that the strength of sexual selection (i) is driven by the relative abundance of males and females prepared to mate (i.e. the operational sex ratio, OSR) and (ii) can be generally estimated by calculating intra‐sexual variation in mating success (e.g. the opportunity for sexual selection, I_s). Here, we demonstrate the problematic nature of these predictions. The OSR and I_s only accurately predict sexual selection under a limited set of circumstances, and more specifically, only when mate monopolization is extremely strong. If mate monopolization is not strong, using OSR or I_s as proxies or measures of sexual selection is expected to produce spurious results that lead to the false conclusion that sexual selection is strong when it is actually weak. These findings call into question the validity of empirical conclusions based on these measures of sexual selection.     

The role of short-tongued insects in floral variation across the range of a style-dimorphic plant

Background and Aims

Heterostyly and related style polymorphisms are suitable model systems to evaluate the importance of functional pollinators in the maintenance of population variability. In Narcissus papyraceus different functional pollinators, incompatibility system and flower morphology have been proposed to influence the maintenance of polymorphism through their effect on disassortative mating. Here a test is done to find out if the visitation rate of long- versus short-tongued pollinators correlates with the morph ratio and if the latter is related to other flower traits of the species across its main geographic range.

Methods

Floral traits from 34 populations in the south-west of the Iberian Peninsula and in north-west Africa were measured, perianth variation was described and a comparison was made of allometric relationships between sex organs and floral tube. Correlations between pollinator guilds, stigma–anther separation of reciprocal morphs (our proxy for disassortative mating) and morph-ratio variation were analysed. Finally, the incompatibility system of the species in the northern and southern borders of its distribution are described.

Key Results

Flowers from southern populations were significantly larger than flowers from centre and northern populations. The abundance of short-styled plants decreased gradually with increasing distance from the core region (the Strait of Gibraltar), with these disappearing only in the northern range. Although there was a significant difference in stigma–anther separation among populations, morph ratio was not associated with reciprocity or floral tube length. Long-style morph frequency increased with short-tongued pollinator visitation rate. Populations from both edges of the distribution range were self-incompatible and within- and between-morph compatible.

Conclusions

The style morph ratio changed gradually, whereas perianth trait variation showed abrupt changes with two morphotypes across the range. The positive relationship between the visitation rate of short-tongued pollinators and the decrease of the short-style morph supports our initial hypothesis. The results highlight the importance of different pollinators in determining the presence of style polymorphism.     

Variation of pollen limitation in the early flowering Mediterranean geophyte Narcissus assoanus (Amaryllidaceae)

A recent literature review indicates that pollen limitation of female fertility is a common feature of flowering plants. Despite the ecological and evolutionary significance of pollen limitation, most studies have only examined fertility in a single population at one time. Here we investigate pollen limitation of fruit and seed set in five populations of Narcissus assoanus, a self- sterile, insect-pollinated geophyte, over 2–3 years in southern France. In common with many early spring flowering plants, pollinator visitation to N. assoanus is often infrequent. Supplemental hand-pollination of flowers with outcross pollen significantly increased overall fruit and seed set by 11% and 19%, respectively. Four of the five populations experienced some pollen limitation during the study. For a given year, there was significant variation in pollen limitation among populations. Two of the populations were pollen limited in one year but not in other years in which they were studied. Seed:ovule ratios for open- and hand-pollinated flowers averaged 0.29 and 0.33, respectively. While hand pollination significantly increased the seed:ovule ratio, the low value obtained indicates that the majority of ovules in flowers do not mature seeds despite hand pollination. The role of genetic and environmental factors governing low seed:ovule ratios in N. assoanus is discussed. Received: 28 December 1999 / Accepted: 6 April 2000     

The population genetics of the B-chromosome polymorphism of Myrmeleotettix maculatus (Orthoptera: Acrididae)

A simple difference equation model of the dynamics of the B-chromosome of Myrmeleotettix maculatus is developed, validated with field data, and analysed. The maintenance of the B in a population is due to a frequency-dependent balance between a net meiotic drive and strong selection against karyotypes containing 2 B's. This equilibrium will be subject to substantial fluctuations even in moderately large populations. In the long run a B is likely to be maintained in a population despite the selection of suppressor genes, which will tend to remain polymorphic in the population. These qualitative conclusions are likely to apply to many B-chromosome systems.     

A measure of sexual selection for interspecific comparisons of species with diversified mating systems and different mortality schedules

Based on variances in components of lifetime reproductive success (LRS), a new index of the intensity of sexual selection was proposed mainly useful for interspecific comparisons. The index was defined as the male to female ratio of the standardized variance in mating efficiency during mating period. The index can to a large extent exclude the effect from natural selection which commonly act on both sexes in mating period and also from mortality in pre-reproductive period. This empirical measure has some defects in the strict sense (i.e., dichotomy of natural and sexual selection), however, it enables interspecific comparisons of the intensity of sexual selection among different taxonomic groups of animals with various mating systems and mortality schedules.     

The influence of environmental quality on sexual selection in Nauphoeta cinerea (Dictyoptera: Blaberidae)

We examined the impact of environmental conditions on the sexpheromone and mating behavior of the cockroach, Nauphoeta cinerea.Previous research on this species has shown that female behaviorduring courtship reflects female mate choice, male behaviorcorrelates with male social status, and the male sex pheromoneis the character used by females to assess males. In the presentstudy, males and females were allowed to develop from adultemergence to sexual maturity in either a high- or low-qualityenvironment. The environment affected the quantities of sexpheromone components. We found significantly less 3-hydroxy-2-butanoneand 4-ethyl-2-methoxyphenol, but not 2-methylthiazolidine, inthe pheromone glands of males from a poor environment. Pheromonequality was also affected; the ratios involving 2-methylthiazolidinewere altered, while the ratio 3-hydroxy-2-butanone to 4-ethyl-2-methoxyphenoldid not change. Development to sexual maturity under these environmentalconditions also influenced male and female sexual behavior.Male courtship activity reflected environmental influences;males from the low-quality environment took longer to initiatecourtship and spent more time copulating with females from allenvironments. Male quality, as assessed by females, was alsoaffected by their environment. Females were slower to respondto the courtship of males from the poor environment, regardlessof the females' own rearing environments. However, females fromthe low-quality environment also took longer to respond to thecourtship, and required more courtship, regardless of the males'rearing environments. Thus, poor environments also increasefemale choosiness. However, there was only one significant interactionterm, suggesting that the environmental effects are generaland that females do not show adaptive plasticity in mate choice.Studies of sexual selection that consider the effects of variableenvironments on behavior as well as the sexually selected morphologyin other systems are likely to provide new insights into thisevolutionary process