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1.  Paternal effects on functional gender account for cryptic dioecy in a perennial plant. 
Natural selection operates on the mating strategies of hermaphrodites through their functional gender, i.e. their relative success as male versus female parents. Because functional gender will tend to be strongly influenced by sex allocation, it is often estimated in plants by counting seeds and pollen grains. However, a plant's functional gender must also depend on the fate of the seeds and pollen grains it produces. We provide clear evidence of a paternal effect on the functional gender of a plant that is independent of the resources invested in pollen. In the Mediterranean tree Fraxinus ornus, males coexist with hermaphrodites that disperse viable pollen and that sire seeds; the population would thus appear to be functionally androdioecious. However, we found that seedlings sired by hermaphrodites grew significantly less well than those sired by males, suggesting that hermaphrodites may be functionally less male than they seem. The observed 1 : 1 sex ratios in F. ornus, which have hitherto been difficult to explain in the light of the seed-siring ability of hermaphrodites, support our interpretation that this species is cryptically dioecious. Our results underscore the importance of considering progeny quality when estimating gender, and caution against inferring androdioecy on the basis of a siring ability of hermaphrodites alone.
PMCID: PMC1691823  PMID: 15451691
2.  Differential costs of reproduction in females and hermaphrodites in a gynodioecious plant 
Annals of Botany  2012;109(6):1159-1164.
Background and Aims
Plants exhibit a variety of reproductive systems where unisexual (females or males) morphs coexist with hermaphrodites. The maintenance of dimorphic and polymorphic reproductive systems may be problematic. For example, to coexist with hermaphrodites the females of gynodioecious species have to compensate for the lack of male function. In our study species, Geranium sylvaticum, a perennial gynodioecious herb, the relative seed fitness advantage of females varies significantly between years within populations as well as among populations. Differences in reproductive investment between females and hermaphrodites may lead to differences in future survival, growth and reproductive success, i.e. to differential costs of reproduction. Since females of this species produce more seeds, higher costs of reproduction in females than in hermaphrodites were expected. Due to the higher costs of reproduction, the yearly variation in reproductive output of females might be more pronounced than that of hermaphrodites.
Using supplemental hand-pollination of females and hermaphrodites of G. sylvaticum we examined if increased reproductive output leads to differential costs of reproduction in terms of survival, probability of flowering, and seed production in the following year.
Key Results
Experimentally increased reproductive output had differential effects on the reproduction of females and hermaphrodites. In hermaphrodites, the probability of flowering decreased significantly in the following year, whereas in females the costs were expressed in terms of decreased future seed production.
When combining the probability of flowering and seed production per plant to estimate the multiplicative change in fitness, female plants showed a 56 % and hermaphrodites showed a 39 % decrease in fitness due to experimentally increased reproduction. Therefore, in total, female plants seem to be more sensitive to the cost of reproduction in terms of seed fitness than hermaphrodites.
PMCID: PMC3336951  PMID: 22419762
Gynodioecy; pollen limitation; hand-pollination; Geranium sylvaticum; cost of reproduction; maintenance of gynodioecy; seed production
3.  Gender plasticity and sexual system stability in Wurmbea 
Annals of Botany  2011;109(3):521-530.
Background and aims
Sexually dimorphic populations are often located in drier habitats than cosexual populations. Gender plasticity (GP), whereby hermaphrodites alter female and male functions depending on resources, and sex-differential plasticity (SDP) between hermaphrodites and unisexuals are predicted to affect sexual system stability. Here, GP and SDP are evaluated in cosexual and gynodioecious Wurmbea biglandulosa and sub-dioecious and dioecious W. dioica.
GP was evaluated under two resource conditions, compared among sexual systems and assessed as to whether (1) males produced perfect flowers and (2) hermaphrodites altered investment in perfect (female function) and total (male function) flowers. SDP was assessed within sexual systems as differences between sex functions of hermaphrodites vs. unisexuals. Males and hermaphrodites were compared to assess whether size thresholds for female function differed among sexual systems. Plasticity costs were evaluated using correlations between female function and male traits in hermaphrodites, and in W. dioica by comparing hermaphrodite and male regressions between plant size and pollen production.
Key Results
In dioecious W. dioica no males exhibited GP, whereas 100 % did in gynodioecious and cosexual W. biglandulosa. In sub-dioecious W. dioica, resources affected GP (high, 66 %; low, 42 %). Hermaphrodites in all sexual systems reduced perfect but not total flowers under low resources. Unisexuals were unaffected, demonstrating SDP for female function only. Thresholds for female function were greater in sub-dioecious W. dioica than in W. biglandulosa. Plasticity costs were detected only in sub-dioecious W. dioica.
SDP for female function could assist female establishment in cosexual populations and maintain females in gynodioecious and sub-dioecious populations. Although the absence of male SDP should stabilize sub-dioecy, plasticity costs would render sub-dioecy unstable, favouring canalized males over hermaphrodites. This study highlights the importance of interactions between environmental conditions and hermaphrodite sex expression for the stability of dimorphic sexual systems.
PMCID: PMC3278287  PMID: 21831858
Dioecy; gender plasticity; gynodioecy; plant sexual systems; sex-allocation plasticity; sex-differential plasticity; sub-dioecy; Wurmbea biglandulosa; Wurmbea dioica
4.  Roots, shoots and reproduction: sexual dimorphism in size and costs of reproductive allocation in an annual herb 
Females tend to be smaller than males in woody dioecious plant species, but they tend to be larger in herbs. The smaller size of females in woody species has been attributed to higher reproductive costs, yet no satisfactory explanation has been provided for their larger size in herbs. Because herbs have higher nitrogen concentrations in their tissues than woody plants, and because pollen is particularly rich in nitrogen, we predicted that male growth would be more compromised by reproduction than female growth. To test this hypothesis, we conducted three experiments on the annual dioecious herb Mercurialis annua. First, we compared the timing of reproduction between males and females and found that males started flowering earlier than females; early flowering is expected to compromise growth more than later flowering. Second, we compared plants allowed to flower with those prevented from flowering by experimental debudding and found that males incurred a higher reproductive cost than females in terms of both biomass and, particularly, nitrogen. Third, we grew plants under varying levels of nitrogen availability and found that although sexual size dimorphism was unaffected by nitrogen, females, but not males, decreased their relative allocation to both roots and reproduction under high nitrogen availability. We propose that males deal with the high cost of pollen production in terms of nitrogen by allocating biomass to nitrogen-harvesting roots, whereas females pay for carbon-rich seeds and fruits by investing in photosynthetic organs. Sexual dimorphism would thus seem to be the outcome of allocation to above- versus below-ground sinks that supply resources (carbon versus nitrogen) limiting the female and male reproduction differentially.
PMCID: PMC2605799  PMID: 18682371
life history; Mercurialis annua; trade-off; nitrogen allocation; carbon allocation
5.  Ecological Context of Breeding System Variation: Sex, Size and Pollination in a (Predominantly) Gynodioecious Shrub 
Annals of Botany  2007;100(7):1547-1556.
Background and Aims
Species that exhibit among-population variation in breeding system are particularly suitable to study the importance of the ecological context for the stability and evolution of gender polymorphism. Geographical variation in breeding system and sex ratio of Daphne laureola (Thymelaeaceae) was examined and their association with environmental conditions, plant and floral display sizes, and pollination environment in a broad geographic scale was analysed.
The proportion of female and hermaphrodite individuals in 38 populations within the Iberian Peninsula was scored. Average local temperature and precipitation from these sites were obtained from interpolation models based on 30 years of data. Pollination success was estimated as stigmatic pollen loads, pollen tubes per ovule and the proportion of unfertilized flowers per individual in a sub-set of hermaphroditic and gynodioecious populations.
Key Results
Daphne laureola is predominantly gynodioecious, but hermaphroditic populations were found in northeastern and southwestern regions, characterized by higher temperatures and lower annual precipitation. In the gynodioecious populations, female plants were larger and bore more flowers than hermaphrodites. However, due to their lower pollination success, females did not consistently produce more seeds than hermaphrodites, which tends to negate a seed production advantage in D. laureola females. In the northeastern hermaphroditic populations, plants were smaller and produced 9–13 times fewer flowers than in the other Iberian regions, and thus presumably had a lower level of geitonogamous self-fertilization. However, in a few southern populations hermaphroditism was not associated with small plant size and low flower production.
The findings highlight that different mechanisms, including abiotic conditions and pollinator service, may account for breeding system variation within a species' distribution range and also suggest that geitonogamy may affect plant breeding system evolution.
PMCID: PMC2759233  PMID: 17933844
Daphne laureola; environmental gradients; floral display; geographic variation; geitonogamy; gynodioecy; pollination success; sex ratio; Thymelaeaceae
6.  Costs of receipt and donation of ejaculates in a simultaneous hermaphrodite 
Sexual conflicts between mating partners can strongly impact the evolutionary trajectories of species. This impact is determined by the balance between the costs and benefits of mating. However, due to sex-specific costs it is unclear how costs compare between males and females. Simultaneous hermaphrodites offer a unique opportunity to determine such costs, since both genders are expressed concurrently. By limiting copulation of focal individuals in pairs of pond snails (Lymnaea stagnalis) to either the male role or the female role, we were able to compare the fecundity of single sex individuals with paired hermaphrodites and non-copulants. Additionally, we examined the investment in sperm and seminal fluid of donors towards feminized snails and hermaphrodites.
Compared to non-mating focal snails, reciprocating individuals as well as male and female copulants experienced a significant fecundity reduction (~40%) after, on average, 3.07 ± 0.12 copulations in their allowed roles (for donors 2.98 ± 0.16 copulations and for recipients 3.14 ± 0.12 copulations). In a single copulation, significantly more sperm was donated to partners that were restricted to mating in the female role than to hermaphrodites, while seminal fluid transfer was unaffected by recipient type.
Our data indicate that the costs of mating in both sex functions are high in L. stagnalis. This conclusion is based on fecundity data collected separately for male and female copulants. Male mating costs result from investment in expensive ejaculates, composed of sperm and seminal fluid. For female copulants, fecundity reduction correlated with transferred sperm numbers in the first copulation, while differences in transferred quantities of seminal fluid were not detected. These findings may point toward a "sperm effect" as a novel feature of pond snail reproductive ecology. In conclusion, sex allocation and sexual conflict both contribute to decreased female fecundity in pond snails.
PMCID: PMC3017047  PMID: 21184682
7.  Genetic differentiation for size at first reproduction through male versus female functions in the widespread Mediterranean tree Pinus pinaster 
Annals of Botany  2012;110(7):1449-1460.
Background and Aims
The study of local adaptation in plant reproductive traits has received substantial attention in short-lived species, but studies conducted on forest trees are scarce. This lack of research on long-lived species represents an important gap in our knowledge, because inferences about selection on the reproduction and life history of short-lived species cannot necessarily be extrapolated to trees. This study considers whether the size for first reproduction is locally adapted across a broad geographical range of the Mediterranean conifer species Pinus pinaster. In particular, the study investigates whether this monoecious species varies genetically among populations in terms of whether individuals start to reproduce through their male function, their female function or both sexual functions simultaneously. Whether differences among populations could be attributed to local adaptation across a climatic gradient is then considered.
Male and female reproduction and growth were measured during early stages of sexual maturity of a P. pinaster common garden comprising 23 populations sampled across the species range. Generalized linear mixed models were used to assess genetic variability of early reproductive life-history traits. Environmental correlations with reproductive life-history traits were tested after controlling for neutral genetic structure provided by 12 nuclear simple sequence repeat markers.
Key Results
Trees tended to reproduce first through their male function, at a size (height) that varied little among source populations. The transition to female reproduction was slower, showed higher levels of variability and was negatively correlated with vegetative growth traits. Several female reproductive traits were correlated with a gradient of growth conditions, even after accounting for neutral genetic structure, with populations from more unfavourable sites tending to commence female reproduction at a lower individual size.
The study represents the first report of genetic variability among populations for differences in the threshold size for first reproduction between male and female sexual functions in a tree species. The relatively uniform size at which individuals begin reproducing through their male function probably represents the fact that pollen dispersal is also relatively invariant among sites. However, the genetic variability in the timing of female reproduction probably reflects environment-dependent costs of cone production. The results also suggest that early sex allocation in this species might evolve under constraints that do not apply to other conifers.
PMCID: PMC3489151  PMID: 23002272
Pinus pinaster; conifers; sex-dependent threshold size for first reproduction; size-dependent sex allocation; clinal variation; neutral genetic structure
8.  Functional characterization of gynodioecy in Fragaria vesca ssp. bracteata (Rosaceae) 
Annals of Botany  2011;109(3):545-552.
Background and Aims
Gynodioecy is a phylogenetically widespread and important sexual system where females coexist with hermaphrodites. Because dioecy can arise from gynodioecy, characterization of gynodioecy in close relatives of dioecious and sub-dioecious species can provide insight into this transition. Thus, we sought to determine whether Fragaria vesca ssp. bracteata, a close relative to F. chiloensis and F. virginiana, exhibits the functional and population genetic hallmarks of a gynodioecious species.
We compared reproductive allocation of females and hermaphrodites grown in the greenhouse and estimated genetic diversity (allelic diversity, heterozygosity) and inbreeding coefficients for field-collected adults of both sexes using simple sequence repeat (SSR) markers. We estimated mating system and early seed fitness from open-pollinated families of both sex morphs.
Key Results
Under greenhouse conditions, females and hermaphrodites allocated similarly to all reproductive traits except flower number, and, as a consequence, females produced 30 % fewer seeds per plant than hermaphrodites. Under natural conditions, hermaphrodites produce seeds by self-fertilization approx. 75 % of the time, and females produced outcrossed seeds with very little biparental inbreeding. Consistent with inbreeding depression, seeds from open-pollinated hermaphrodites were less likely to germinate than those from females, and family-level estimates of hermaphrodite selfing rates were negatively correlated with germination success and speed. Furthermore, estimates of inbreeding depression based on genetic markers and population genetic theory indicate that inbreeding depression in the field could be high.
The joint consideration of allocation and mating system suggests that compensation may be sufficient to maintain females given the current understanding of sex determination. Fragaria vesca ssp. bracteata exhibited similar sex morph-dependent patterns of mating system and genetic diversity, but less reproductive trait dimorphism, than its sub-dioecious and dioecious congeners.
PMCID: PMC3278295  PMID: 22052984
Dioecy; Fragaria chiloensis; Fragaria vesca ssp. bracteata; Fragaria virginiana; gynodioecy; inbreeding depression; selfing rate; SSR; strawberry
9.  Hermaphrodite life history and the maintenance of partial selfing in experimental populations of Caenorhabditis elegans 
Classic population genetics theory predicts that mixed reproductive systems, where self reproduction (selfing) and outcrossing co-exist, should not be as common as they are in nature. One means of reconciling theory with observations is to recognize that sexual conflict between males and hermaphrodites and/or constraints in the allocation of resources towards sex functions in hermaphrodites can balance the fitness components of selfing and outcrossing.
Using experimental evolution in Caenorhabditis elegans, we test whether the adaptive maintenance of partial selfing is due to sexual conflict and/or to the evolution of sex allocation towards male function in hermaphrodites. For this, we characterized the reproductive schedule and longevity patterns in hermaphrodites under selfing and under outcrossing with naïve males that did not have the opportunity to evolve with them. A shift in reproductive schedule towards earlier reproduction would be indicative of adaptation in our imposed life-cycle, while longevity is expected to evolve as a response to the harm that males impinge on hermaphrodites upon mating. To determine adaptation in the absence of constraints in sex allocation, we also characterized the life history of females that reproduced during experimental evolution through obligate mating with males. As expected with adaptation, we find that after 100 generations of experimental evolution, selfing hermaphrodites and females showed improved reproduction at earlier ages. We did not observe similar reproductive shifts in outcrossed hermaphrodites. We further find increased longevity in outcrossed females after evolution but not in outcrossed hermaphrodites, a result that indicates that sexual conflicts were likely more prevalent under male–female evolution than under male-hermaphrodite evolution.
Taken together, our findings suggest that the adaptive maintenance of partial selfing during C. elegans experimental evolution resulted from the evolution of sex allocation towards male function in hermaphrodites.
PMCID: PMC4052797  PMID: 24891140
10.  Chemical and Physical Defense Traits in Two Sexual Forms of Opuntia robusta in Central Eastern Mexico 
PLoS ONE  2014;9(3):e89535.
Sexually dimorphic plants provide an excellent opportunity for examining the differences in the extent of their defense against herbivores because they exhibit sex-related differences in reproductive investment. Such differences enable comparison of the sex with high reproduction expenses with the sex that expends less. The more costly sex is usually also better defended against herbivores. Generally, females are considered more valuable than hermaphrodites in terms of fitness; however, hermaphrodites are more valuable if they can produce seed by autonomous selfing, provided that the inbreeding depression is low and pollen is limited. We studied a gynodioecious population of Opuntia robusta from Central-Eastern Mexico, which has been reported to be trioecious, dioecious, or hermaphrodite, and addressed the following questions: 1) Is the hermaphrodite's reproductive output higher than the female's, and are hermaphrodites thus better defended? 2) Are plant tissues differentially defended? 3) Do trade-offs exist among different physical defense traits? and 4) among physical and chemical defense traits? We found that 1) hermaphrodites had a higher seed output and more spines per areola than females and that their spines contained less moisture. Non-reproductive hermaphrodite cladodes contained more total phenolic compounds (TPCs) than female ones. In addition, 2) hermaphrodite reproductive cladodes bore more spines than female cladodes, and 3) and 4) we found a negative relationship between spine number per areola and areola number per cladode and a positive relationship between spine number per areola per plant and TPC concentration per plant. Non-reproductive hermaphrodite cladodes contained a higher concentration of TPCs than female cladodes, and parental cladodes contained fewer TPCs than both reproductive and empty cladodes.
PMCID: PMC3943789  PMID: 24599143
11.  The Phylogeny and Biogeographic History of Ashes (Fraxinus, Oleaceae) Highlight the Roles of Migration and Vicariance in the Diversification of Temperate Trees 
PLoS ONE  2013;8(11):e80431.
The cosmopolitan genus Fraxinus, which comprises about 40 species of temperate trees and shrubs occupying various habitats in the Northern Hemisphere, represents a useful model to study speciation in long-lived angiosperms. We used nuclear external transcribed spacers (nETS), phantastica gene sequences, and two chloroplast loci (trnH-psbA and rpl32-trnL) in combination with previously published and newly obtained nITS sequences to produce a time-calibrated multi-locus phylogeny of the genus. We then inferred the biogeographic history and evolution of floral morphology. An early dispersal event could be inferred from North America to Asia during the Oligocene, leading to the diversification of the section Melioides sensus lato. Another intercontinental dispersal originating from the Eurasian section of Fraxinus could be dated from the Miocene and resulted in the speciation of F. nigra in North America. In addition, vicariance was inferred to account for the distribution of the other Old World species (sections Sciadanthus, Fraxinus and Ornus). Geographic speciation likely involving dispersal and vicariance could also be inferred from the phylogenetic grouping of geographically close taxa. Molecular dating suggested that the initial divergence of the taxonomical sections occurred during the middle and late Eocene and Oligocene periods, whereas diversification within sections occurred mostly during the late Oligocene and Miocene, which is consistent with the climate warming and accompanying large distributional changes observed during these periods. These various results underline the importance of dispersal and vicariance in promoting geographic speciation and diversification in Fraxinus. Similarities in life history, reproductive and demographic attributes as well as geographical distribution patterns suggest that many other temperate trees should exhibit similar speciation patterns. On the other hand, the observed parallel evolution and reversions in floral morphology would imply a major influence of environmental pressure. The phylogeny obtained and its biogeographical implications should facilitate future studies on the evolution of complex adaptive characters, such as habitat preference, and their possible roles in promoting divergent evolution in trees.
PMCID: PMC3837005  PMID: 24278282
12.  Sexual dimorphism in a dioecious population of the wind-pollinated herb Mercurialis annua: the interactive effects of resource availability and competition 
Annals of Botany  2011;107(6):1039-1045.
Background and Aims
Male-biased sex allocation commonly occurs in wind-pollinated hermaphroditic plants, and is often positively associated with size, notably in terms of height. Currently, it is not well established whether a corresponding pattern holds for dioecious plants: do males of wind-pollinated species exhibit greater reproductive allocation than females? Here, sexual dimorphism is investigated in terms of life history trade-offs in a dioecious population of the wind-pollinated ruderal herb Mercurialis annua.
The allocation strategies of males and females grown under different soil nutrient availability and competitive (i.e. no, male or female competitor) regimes were compared.
Key Results
Male reproductive allocation increased disproportionately with biomass, and was greater than that of females when grown in rich soils. Sexual morphs differentially adjusted their reproductive allocation in response to local environmental conditions. In particular, males reduced their reproductive allocation in poor soils, whereas females increased theirs, especially when competing with another female rather than growing alone. Finally, males displayed smaller above-ground vegetative sizes than females, but neither nutrient availability nor competition had a strong independent effect on relative size disparities between the sexes.
Selection appears to favour plasticity in reproductive allocation in dioecious M. annua, thereby maintaining a relatively constant size hierarchy between sexual morphs. In common with other dioecious species, there seems to be little divergence in the niches occupied by males and females of M. annua.
PMCID: PMC3080628  PMID: 21385775
Life history trade-offs; competition; wind pollination; separate sexes; sex allocation; sexual size dimorphism; reproductive allocation; resource availability; Mercurialis annua
13.  Flower Bud Abortion Influences Clonal Growth and Sexual Dimorphism in the Understorey Dioecious Shrub Aucuba japonica (Cornaceae) 
Annals of Botany  2002;89(6):675-681.
Sexual differences were investigated to determine the significance of flower bud abortion in the dioecious shrub Aucuba japonica Thunb. The mean number of flowers per inflorescence and the mean number of flowering inflorescences (as opposed to aborted inflorescences) per individual were greater in males than in females in 1997 and 1998. Reproductive investment by males was 0·4‐times (1997) and 1·4‐times (1998) that by females. In addition, females aborted 30·9 % (1997) and 42·7 % (1998) of their total flower buds without blooming, whereas no male flower buds aborted. One of the architectural traits of this shrub is that in the year that a flower bud is produced at the shoot apex, the shoot will branch into two or more shoots. Thus, there was less sexual difference in the number of current shoots per individual than there was in the number of flowering inflorescences. The relationship between annual growth and reproduction, and the probability of reproduction in the following year, suggested that the higher investment in female reproduction was manifested as a cost for reproductive frequency rather than as a cost for annual growth. The spatial distribution of both males and females was clumped, which may be the result of clonal growth. In addition, overall sex ratios were not skewed and the number of sprouts did not differ significantly between sexes. These results suggested that flower bud abortion by females might reduce sexual dimorphism in terms of clonal growth.
PMCID: PMC4233831  PMID: 12102522
Aucuba japonica; clonal growth; flower bud abortion; forest understorey; branching; reproductive investment; sexual dimorphism; sprout
14.  Experimental defoliation affects male but not female reproductive performance of the tropical monoecious plant Croton suberosus (Euphorbiaceae) 
Annals of Botany  2010;106(2):359-369.
Background and Aims
Monoecious plants have the capacity to allocate resources separately to male and female functions more easily than hermaphrodites. This can be advantageous against environmental stresses such as leaf herbivory. However, studies showing effects of herbivory on male and female functions and on the interaction with the plant's pollinators are limited, particularly in tropical plants. Here, the effects of experimental defoliation were examined in the monoecious shrub Croton suberosus (Euphorbiaceae), a wasp-pollinated species from a Mexican tropical dry forest.
Three defoliation treatments were applied: 0 % (control), 25 % (low) or 75 % (high) of plant leaf area removed. Vegetative (production of new leaves) and reproductive (pistillate and staminate flower production, pollen viability, nectar production, fruit set, and seed set) performance variables, and the abundance and activity of floral visitors were examined.
Key Results
Defoliated plants overcompensated for tissue loss by producing more new leaves than control plants. Production of staminate flowers gradually decreased with increasing defoliation and the floral sex ratio (staminate : pistillate flowers) was drastically reduced in high-defoliation plants. In contrast, female reproductive performance (pistillate flower production, fruit set and seed set) and pollinator visitation and abundance were not impacted by defoliation.
The asymmetrical effects of defoliation on male and female traits of C. suberosus may be due to the temporal and spatial flexibility in the allocation of resources deployed by monoecious plants. We posit that this helps to maintain the plant's pollination success in the face of leaf herbivory stress.
PMCID: PMC2908170  PMID: 20519239
Euphorbiaceae; floral sex ratio; foliar herbivory; leaf production; nectar production; Neotropical dry forest; plant–insect interactions; pollen production; pollination success; pollinator activity
15.  Condition Dependent Effects on Sex Allocation and Reproductive Effort in Sequential Hermaphrodites 
PLoS ONE  2014;9(10):e109626.
Theory predicts the optimal timing of sex change will be the age or size at which half of an individual's expected fitness comes through reproduction as a male and half through reproduction as a female. In this way, sex allocation across the lifetime of a sequential hermaphrodite parallels the sex allocation of an outbreeding species exhibiting a 1∶1 ratio of sons to daughters. However, the expectation of a 1∶1 sex ratio is sensitive to variation in individual condition. If individuals within a population vary in condition, high-condition individuals are predicted to make increased allocations to the sex with the higher variance in reproductive success. An oft-cited example of this effect is seen in red deer, Cervus elaphus, in which mothers of high condition are more likely to produce sons, while those in low condition are more likely to produce daughters. Here, we show that individual condition is predicted to similarly affect the pattern of sex allocation, and thus the allocation of reproductive effort, in sequential hermaphrodites. High-condition sex-changers are expected to obtain more than half of their fitness in the high-payoff second sex and, as a result, are expected to reduce the allocation of reproductive effort in the initial sex. While the sex ratio in populations of sequential hermaphrodites is always skewed towards an excess of the initial sex, condition dependence is predicted to increase this effect.
PMCID: PMC4193790  PMID: 25302941
16.  Gender-specific costs of reproduction on vegetative growth and physiological performance in the dioecious shrub Corema album 
Annals of Botany  2010;106(6):989-998.
Background and Aims
Reproductive costs imply trade-offs in resource distribution at the physiological level, expressed as changes in future growth and/or reproduction. In dioecious species, females generally endure higher reproductive effort, although this is not necessarily expressed through higher somatic costs, as compensatory mechanisms may foster resource uptake during reproduction.
To assess effects of reproductive allocation on vegetative growth and physiological response in terms of costs and compensation mechanisms, a manipulative experiment of inflorescence bud removal was carried out in the sexually dimorphic species Corema album. Over two consecutive growing seasons, vegetative growth patterns, water status and photochemical efficiency were measured to evaluate gender-related differences.
Key Results
Suppression of reproductive allocation resulted in a direct reduction in somatic costs of reproduction, expressed through changes in growth variables and plant physiological status. Inflorescence bud removal was related to an increase in shoot elongation and water potential in male and female plants. The response to inflorescence bud removal showed gender-related differences that were related to the moment of maximum reproductive effort in each sexual form: flowering in males and fruiting in females. Delayed costs of reproduction were found in both water status and growth variables, showing gender-related differences in resource storage and use.
Results are consistent with the existence of a trade-off between reproductive and vegetative biomass, indicating that reproduction and growth depend on the same resource pool. Gender-related morphological and physiological differences arise as a response to different reproductive resource requirements. Delayed somatic costs provide evidence of gender-related differences in resource allocation and storage. Adaptive differences between genders in C. album may arise through the development of mechanisms which compensate for the cost of reproduction.
PMCID: PMC2990667  PMID: 20884627
Corema album; costs of reproduction; flower removal; photochemical efficiency; sexual dimorphism; trade-off; water potential
17.  Between-year Changes in Functional Gender Expression of Palicourea padifolia (Rubiaceae), a Distylous, Hummingbird-pollinated Shrub 
Annals of Botany  2004;95(2):371-378.
• Background and Aims Because distylous species have two hermaphroditic style-length floral morphs, they face two sex allocation problems: the equilibrium morph ratio and the optimal allocation to pollen and seed production in each floral morph. Gender specialization is expected among distylous species when floral morphs differ in reproductive output. However, spatio-temporal variability in female reproductive output between morphs needs to be investigated to fully understand patterns of sexual expression and gender specialization in distylous plants. Between-year variation in flower and fruit production of hummingbird-pollinated Palicourea padifolia (Rubiaceae) was examined, focusing on functional gender expression of long- and short-styled morphs and comparing their reproductive performance in five consecutive years (1998–2002).
• Methods Between-year variation in inflorescence, floral bud and fruit production was monitored and quantified. These traits were then used as parameters to determine functional gender differences between floral morphs through time.
• Key Results Inflorescence production varied among years but no significant differences were found between floral morphs. Long-styled plants initiated more floral buds per inflorescence every year than short-styled plants, suggesting higher allocation to pollinator attraction and, potentially, an increase in male fitness through pollen donation. Although fruit production was similar between morphs, their functional gender shifted across years.
• Conclusions The gender expression inconsistency across years is surprising because a number of floral characters and attributes that contribute to differently attracting and rewarding effective pollinators in P. padifolia suggest gender specialization. The evidence that morphs of distylous species might specialize in functional gender mostly comes from differences among populations in seed production and non-equilibrium morph ratios based on 1-year field population surveys. The results suggest that more sampling through time is needed to detect gender specialization among distylous species with a perennial habit.
PMCID: PMC4246829  PMID: 15546929
Cloud forest; distyly; gender expression; hummingbirds; Palicourea padifolia; Rubiaceae; sex-allocation theory
18.  Floral dimorphism in plant populations with combined versus separate sexes 
Annals of Botany  2011;108(4):765-776.
Background and Aims
Dimorphism among floral traits can evolve through variation in selection intensity between female and male performance, especially when sex functions are separated between flowers on a plant (monoecy), or between individuals (dioecy). In animal-pollinated species, male floral traits are predicted to be larger because competition for pollinators should favour larger displays. Floral dimorphism may be greater in dioecious than monoecious populations because of trade-offs between female and male function and opportunities for selfing in hermaphrodites.
These predictions were tested by surveying flower size, total flowers per inflorescence and daily display size in the insect-pollinated Sagittaria latifolia (Alismataceae). This species is useful for comparative analysis because populations are mostly either monoecious or dioecious. We examined floral dimorphism in 13 monoecious and 16 dioecious populations in eastern North America.
Key Results
Male flowers were significantly larger than female flowers in monoecious and dioecious populations, but there was no evidence for greater flower size dimorphism in dioecious populations despite their larger flower sizes overall. Although inflorescences in both dioecious and monoecious populations produced more male flowers, daily floral displays were significantly larger for female than male function due to more synchronous female flower opening. Daily floral display dimorphism was significantly greater in dioecious populations, due to greater female daily floral displays. There was a positive relationship between mean flower size and total flowers per inflorescence for both sexes in dioecious populations, but no relationship for either sex function in monoecious populations. Flower size dimorphism was positively correlated with the frequencies of females in dioecious populations.
The increased size and number of male flowers and protracted male floral displays in S. latifolia are probably shaped by sexual selection for more effective pollen dispersal.
PMCID: PMC3170149  PMID: 21385777
Sexual dimorphism; flower size; daily floral display; sexual selection; sex ratios; monoecy; dioecy; Sagittaria latifolia
19.  Two's Company, Three's a Crowd: Experimental Evaluation of the Evolutionary Maintenance of Trioecy in Mercurialis annua (Euphorbiaceae) 
PLoS ONE  2012;7(4):e35597.
Trioecy is an uncommon sexual system in which males, females, and hermaphrodites co-occur as three clearly different gender classes. The evolutionary stability of trioecy is unclear, but would depend on factors such as hermaphroditic sex allocation and rates of outcrossing vs. selfing. Here, trioecious populations of Mercurialis annua are described for the first time. We examined the frequencies of females, males and hermaphrodites across ten natural populations and evaluated the association between the frequency of females and plant densities. Previous studies have shown that selfing rates in this species are density-dependent and are reduced in the presence of males, which produce substantially more pollen than hermaphrodites. Accordingly, we examined the evolutionary stability of trioecy using an experiment in which we (a) indirectly manipulated selfing rates by altering plant densities and the frequency of males in a fully factorial manner across 20 experimental plots and (b) examined the effect of these manipulations on the frequency of the three sex phenotypes in the next generation of plants. In the parental generation, we measured the seed and pollen allocations of hermaphrodites and compared them with allocations by unisexual plants. In natural populations, females occurred at higher frequencies in denser patches, a finding consistent with our expectations. Under our experimental conditions, however, no combination of plant densities and male frequencies was associated with increased frequencies of females. Our results suggest that the factors that regulate female frequencies in trioecious populations of M. annua are independent of those regulating male frequencies (density), and that the stable co-existence of all three sex phenotypes within populations is unlikely.
PMCID: PMC3330815  PMID: 22532862
20.  Soil water content and patterns of allocation to below- and above-ground biomass in the sexes of the subdioecious plant Honckenya peploides 
Annals of Botany  2012;110(4):839-848.
Background and aims
Dioecious plants often show sex-specific differences in growth and biomass allocation. These differences have been explained as a consequence of the different reproductive functions performed by the sexes. Empirical evidence strongly supports a greater reproductive investment in females. Sex differences in allocation may determine the performance of each sex in different habitats and therefore might explain the spatial segregation of the sexes described in many dimorphic plants. Here, an investigation was made of the sexual dimorphism in seasonal patterns of biomass allocation in the subdioecious perennial herb Honckenya peploides, a species that grows in embryo dunes (i.e. the youngest coastal dune formation) and displays spatial segregation of the sexes at the studied site. The water content in the soil of the male- and female-plant habitats at different times throughout the season was also examined.
The seasonal patterns of soil-water availability and biomass allocation were compared in two consecutive years in male and female H. peploides plants by collecting soil and plant samples in natural populations. Vertical profiles of below-ground biomass and water content were studied by sampling soil in male- and female-plant habitats at different soil depths.
Key Results
The sexes of H. peploides differed in their seasonal patterns of biomass allocation to reproduction. Males invested twice as much in reproduction than females early in the season, but sexual differences became reversed as the season progressed. No differences were found in above-ground biomass between the sexes, but the allocation of biomass to below-ground structures varied differently in depth for males and females, with females usually having greater below-ground biomass than males. In addition, male and female plants of H. peploides had different water-content profiles in the soil where they were growing and, when differences existed (usually in the upper layers of the soil), the water content of the soil was higher for the female plants had than for the male plants.
Sex-differential timing of investment in reproduction and differential availability and use of resources from the soil (particularly water) are factors that probably offset the costs of reproduction in the above-ground growth in males and females of H. peploides. The results suggest that the patterns of spatial segregation of the sexes observed in H. peploides may contribute to maximize each sex's growth and reproduction.
PMCID: PMC3423814  PMID: 22782243
Dioecy; biomass allocation; below-ground structures; reproductive effort; spatial segregation; water availability
21.  When, How and How Much: Gender-specific Resource-use Strategies in the Dioecious Tree Juniperus thurifera  
Annals of Botany  2006;98(4):885-889.
• Background and Aims In dioecious species male and female plants experience different selective pressures and often incur different reproductive costs. An increase in reproductive investment habitually results in a reduction of the resources available to other demands, such as vegetative growth. Tree-ring growth is an integrative measure that tracks vegetative investment through the plant's entire life span. This allows the study of gender-specific vegetative allocation strategies in dioecious tree species thoughout their life stages.
• Methods Standard dendrochronological procedures were used to measure tree-ring width. Analyses of time-series were made by means of General Mixed Models with correction of autocorrelated values by the use of an autoregressive covariance structure of order one. Bootstrapped correlation functions were used to study the relationship between climate and tree-ring width.
• Key Results Male and female trees invest a similar amount of resources to ring growth during the early life stages of Juniperus thurifera. However, after reaching sexual maturity, tree-ring growth is reduced for both sexes. Furthermore, females experience a significantly stronger reduction in growth than males, which indicates a lower vegetative allocation in females. In addition, growth was positively correlated with precipitation from the current winter and spring in male trees but only to current spring precipitation in females.
• Conclusions Once sexual maturity is achieved, tree rings grow proportionally more in males than in females. Differences in tree-ring growth between the genders could be a strategy to respond to different reproductive demands. Therefore, and responding to the questions of when, how and how much asked in the title, it is shown that male trees invest more resources to growth than female trees only after reaching sexual maturity, and they use these resources in a different temporal way.
PMCID: PMC2806176  PMID: 16905569
Dendrochronology; sexual maturity; resource allocation; trade-offs; Juniperus
22.  Sex Allocation of Females and Hermaphrodites in the Gynodioecious Geranium sylvaticum 
Annals of Botany  2003;92(2):207-213.
Seed production and patterns of sex allocation were studied in female and hermaphroditic plants in two gynodioecious populations of Geranium sylvaticum (Geraniaceae). Females produced more flower buds and seeds than hermaphrodites in one of the two study populations. The other female traits measured (pistil biomass, seed number per fruit, individual seed mass) did not differ between the gender morphs. The relative seed fitness of hermaphrodites differed between the study populations, with hermaphrodites gaining less of their fitness through female function in the population with a high frequency of females. However, the amount and size of pollen produced by hermaphrodites did not differ between populations. The number of flower buds was positively correlated with seed production in females, whereas in hermaphrodites a positive correlation between number of buds and seed production was found in only one of the two study populations. These results suggest that fitness gain through female function is labile in hermaphrodites of this species, and is probably affected by environmental factors such as the sex ratio of the population.
PMCID: PMC4243648  PMID: 12814954
Geranium sylvaticum; Geraniaceae; gynodioecy; sex allocation; pollen production
23.  Preferences of Pollinators and Herbivores in Gynodioecious Geranium sylvaticum 
Annals of Botany  2005;95(5):879-886.
• Background and Aims For the maintenance of gynodioecy (i.e. the coexistence of female and hermaphroditic plants), females need to compensate for the lack of pollen production through higher seed production or better progeny quality compared to hermaphrodites. In Geranium sylvaticum, females produce more seeds per flower than hermaphrodites. This difference in seed production might be modified by biological interactions with pollinators and herbivores that may favour one sex and thus affect the maintenance of gynodioecy.
• Methods Sexual dimorphism in flower size and flowering phenology, and in attractiveness to pollinators, pre-dispersal seed predators and floral herbivores were examined in natural populations of G. sylvaticum.
• Key Results Pollinators preferred hermaphrodites 25 % more often than females in two of the three study populations, and floral herbivores attacked hermaphrodites 15 % more often than females in two of the six study populations. These preferences might be explained by the larger flower size of hermaphrodites. In contrast, seed predators did not prefer either sex.
• Conclusions The data suggest that pollinator preference does not benefit females, whereas the higher floral herbivory of hermaphrodites might enhance the maintenance of females in G. sylvaticum. Thus, although the data support the view that ecological factors may contribute to the maintenance of gynodioecy, they also suggest that these contributions may vary across populations and that they may function in opposite directions.
PMCID: PMC4246743  PMID: 15705604
Floral herbivory; flowering phenology; Geraniaceae; Geranium sylvaticum; gynodioecy; petal size; pollinator attraction; seed predation
24.  Maternal investment in relation to sex ratio and offspring number in a small mammal – a case for Trivers and Willard theory? 
The Journal of Animal Ecology  2009;78(5):1007-1014.
Optimal parental sex allocation depends on the balance between the costs of investing into sons vs. daughters and the benefits calculated as fitness returns. The outcome of this equation varies with the life history of the species, as well as the state of the individual and the quality of the environment.We studied maternal allocation and subsequent fecundity costs of bank voles, Myodes glareolus, by manipulating both the postnatal sex ratio (all-male/all-female litters) and the quality of rearing environment (through manipulation of litter size by −2/+2 pups) of their offspring in a laboratory setting.We found that mothers clearly biased their allocation to female rather than male offspring regardless of their own body condition. Male pups had a significantly lower growth rate than female pups, so that at weaning, males from enlarged litters were the smallest. Mothers produced more milk for female litters and also defended them more intensively than male offspring.The results agree with the predictions based on the bank vole life history: there will be selection for greater investment in daughters rather than sons, as a larger size seems to be more influencial for female reproductive success in this species. Our finding could be a general rule in highly polygynous, but weakly dimorphic small mammals where females are territorial.The results disagree with the narrow sense Trivers & Willard hypothesis, which states that in polygynous mammals that show higher variation in male than in female reproductive success, high-quality mothers are expected to invest more in sons than in daughters.
PMCID: PMC2779470  PMID: 19549147
cost of reproduction; litter size manipulation; nest defence; polygynous mating system; sexual size dimorphism
25.  The link between immunity and life history traits in scleractinian corals 
PeerJ  2014;2:e628.
Immunity is an important biological trait that influences the survival of individuals and the fitness of a species. Immune defenses are costly and likely compete for energy with other life-history traits, such as reproduction and growth, affecting the overall fitness of a species. Competition among these traits in scleractinian corals could influence the dynamics and structural integrity of coral reef communities. Due to variability in biological traits within populations and across species, it is likely that coral colonies within population/species adjust their immune system to the available resources. In corals, the innate immune system is composed of various pathways. The immune system components can be assessed in the absence (constitutive levels) and/or presence of stressors/pathogens (immune response). Comparisons of the constitutive levels of three immune pathways (melanin synthesis, antioxidant and antimicrobial) of closely related species of Scleractinian corals allowed to determine the link between immunity and reproduction and colony growth. First, we explored differences in constitutive immunity among closely related coral species of the genus Meandrina with different reproductive patterns (gonochoric vs. hermaphrodite). We then compared fast-growing branching vs. slow-growing massive Porites to test co-variation between constitutive immunity and growth rates and morphology in corals. Results indicate that there seems to be a relationship between constitutive immunity and sexual pattern with gonochoric species showing significantly higher levels of immunity than hermaphrodites. Therefore, gonochoric species maybe better suited to resist infections and overcome stressors. Constitutive immunity varied in relation with growth rates and colony morphology, but each species showed contrasting trends within the studied immune pathways. Fast-growing branching species appear to invest more in relatively low cost pathways of the immune system than slow-growing massive species. In corals, energetic investments in life-history traits such as reproduction and growth rate (higher energy investment) seem to have a significant impact on their capacity to respond to stressors, including infectious diseases and coral bleaching. These differences in energy investment are critical in the light of the recent environmental challenges linked to global climate change affecting these organisms. Understanding physiological trade-offs, especially those involving the immune system, will improve our understanding as to how corals could/will respond and survive in future adverse environmental conditions associated with climate change.
PMCID: PMC4217183  PMID: 25374778
Coral disease; Constitutive and innate immunity; Hermaphrodite; Gonochoric; Colony morphology; Trade-off; Resource allocation; Scleractinia; Caribbean corals; Biological traits

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