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Sex allocation

Sex allocation refers to the partitioning of an organism's reproductive resources between the production of male and female offspring or, in hermaphroditic species, between male and female reproductive functions, with the goal of maximizing evolutionary fitness.^[1] This concept is central to evolutionary biology, where sex allocation theory models how natural selection shapes these investment decisions across diverse taxa, including animals, plants, and fungi.^[2] Key aspects include the proportion of resources devoted to each sex—often measured as the sex ratio—and adjustments in response to ecological, genetic, or environmental factors.^[3] A foundational prediction of sex allocation theory is Fisher's principle, which posits that, under most conditions, parents should invest equally in sons and daughters, leading to a population sex ratio of 1:1.^[3] This equilibrium arises from frequency-dependent selection: when one sex becomes rare, individuals producing more of that sex gain a fitness advantage, stabilizing the ratio as total parental expenditure balances across sexes.^[2] Fisher's principle, originally articulated in 1930, explains the prevalence of equal sex ratios in many species and serves as the baseline for understanding deviations.^[4] Deviations from equality occur when fitness returns differ between sexes, such as in cases of local mate competition, where offspring mate locally, favoring female-biased ratios to reduce brother-brother competition.^[2] The Trivers-Willard hypothesis predicts condition-dependent biases, with parents in good condition producing more of the sex with higher reproductive variance (often males in polygynous mammals), while those in poor condition favor the opposite.^[2] In haplodiploid insects like bees, genetic systems lead to female-biased allocation due to asymmetric relatedness, often resulting in sexual conflicts over control.^[1] These models highlight how sex allocation integrates with mating systems, parental investment, and environmental cues, influencing phenomena like sequential hermaphroditism and brood sex ratio adjustments.^[5]

Fundamental Theory

Fisher's Principle

In panmictic populations—where mating occurs randomly—Ronald Fisher provided the foundational explanation for why parents typically invest equally in sons and daughters. In his seminal 1930 book, The Genetical Theory of Natural Selection, Fisher argued that the sex ratio evolves to an equilibrium where parental expenditure on male and female offspring is equalized. This equilibrium arises due to frequency-dependent selection: when one sex becomes relatively rare, individuals of that sex experience higher mating success and thus greater reproductive returns per unit of parental investment, favoring the production of the rarer sex until balance is restored. Fisher's insight demonstrated that any deviation from equality would be unstable, as parents producing the underrepresented sex would have a fitness advantage, driving the population sex ratio back toward 1:1.^[6] The mathematical foundation of Fisher's principle emphasizes equal investment in the two sexes. Let r denote the proportion of total investment allocated to males, with $1 - r to females. At equilibrium, the marginal fitness returns from investing in either sex are equal, leading to r = 0.5 regardless of the relative costs C_m (cost per male offspring) and C_f (cost per female). The numerical sex ratio adjusts accordingly, with the ratio of numbers produced N_m / N_f = C_f / C_m, ensuring equal total investment in each sex. For example, if males cost twice as much as females (C_m = 2 C_f), parents should produce twice as many females as males numerically to balance investment.^[6] To see the frequency dependence explicitly, consider a population where the proportion of males is p. In the standard demographic model, the relative reproductive success per male offspring is proportional to $1/p (or more precisely (1-p)/p, due to higher mating opportunities when males are rare), while that per female offspring is constant (each female has fixed reproductive output independent of the number of females). Parental fitness is then maximized when investment balances these returns, stabilizing at equal investment (p \approx 0.5 when costs are equal). An excess of one sex reduces its per capita reproductive success, creating selection pressure for parents to produce more of the opposite sex.^[6] This principle is exemplified in many outcrossing animal species without complicating social or genetic factors, such as birds and mammals, where observed primary sex ratios at birth or hatching closely approximate 1:1. For instance, in humans and domestic chickens, large-scale data confirm near-equal numbers of male and female offspring under random mating conditions.

Parental Investment and Equal Allocation

Parental investment refers to the total resources that parents allocate to offspring production and care, including gametes, nutrients, and post-natal provisioning, which are divided between male and female progeny.^[7] In the context of sex allocation, this investment determines the relative costs of producing sons versus daughters, influencing the evolution of sex ratios to maximize parental fitness. Under the equilibrium condition derived from frequency-dependent selection, parents that disproportionately invest in the cheaper sex face invasion by mutants that favor the rarer, more valuable sex, leading to equal total investment in both sexes across the population. This logic builds on the core frequency-dependent mechanism where the fitness returns to producing each sex equalize at equilibrium. Robert Trivers refined this in 1972 by emphasizing that equality applies to investment in terms of reproductive value, not merely numerical proportions; for instance, if males require greater resources due to larger size or higher care demands, parents should produce fewer males to achieve parity in total fitness returns from each sex.^[7] Empirical observations in birds and mammals support this equal investment model, with primary sex ratios (at fertilization or hatching/birth) typically close to 1:1 when costs are similar between sexes.^[8] In many avian species, such as passerines, hatching sex ratios approximate 50:50, aligning with equal resource allocation under uniform conditions.^[8] Similarly, in mammals like red deer, birth sex ratios hover near 1:1, reflecting balanced parental investment despite minor male biases attributable to slight cost differences. Primary and secondary sex ratios (at adulthood) often align to maintain this equilibrium, as post-natal mortality differences are offset by adjustments in initial allocation to ensure equal lifetime investment. Deviations from 1:1 occur primarily when production costs differ significantly or mating patterns are non-random, though these represent exceptions to the baseline equality.

Local Mate Competition

Local mate competition (LMC) arises in structured populations where mating occurs primarily among siblings within discrete patches, such as hosts or refuges, leading mothers to evolve female-biased offspring sex ratios to maximize inclusive fitness.^[9] This contrasts with Fisher's principle, which predicts equal investment in sons and daughters under random mating in large, panmictic populations.^[9] In LMC scenarios, brothers compete intensely for local mates, diminishing the marginal fitness returns of additional sons relative to daughters, who disperse to new patches.^[9] W.D. Hamilton's 1967 model formalizes LMC by assuming discrete mating groups founded by a small number of inseminated females (foundresses), random mating within groups, and no inter-group mating, with females controlling offspring sex via fertilized (female) or unfertilized (male) eggs in haplodiploid systems.^[9] The key prediction is that when few foundresses colonize a patch—such as one or two—mothers should produce highly female-biased broods, as a single male can fertilize all sisters, while excess males gain little from local competition.^[9] For instance, in haplodiploid insects like fig wasps, where females oviposit into fig syconia that serve as isolated mating arenas, solitary foundresses produce broods with ratios approaching all daughters.^[10] The optimal proportion of male offspring r in Hamilton's model is given by

r = \frac{n-1}{2n},

where n is the number of foundresses per patch; this yields r \to 0 as n \to 1 (extreme female bias) and approaches 0.5 as n \to \infty (equal sex ratio under global mating).^[9] This formula derives from equating the inclusive fitness gains from sons and daughters, accounting for local competition that reduces male mating success.^[9] Inclusive fitness benefits drive the female bias, particularly in haplodiploids, where a female is related to her sisters by 0.75 (three-quarters) but to her brothers by only 0.25 (one-quarter), making daughters more valuable transmitters of her genes than sons under LMC.^[9] Males, related equally to brothers and sisters (0.5), favor equal ratios but lack control over sex allocation.^[9] This asymmetry amplifies selection for female-biased strategies when sibling mating is clumped. Empirical support is strong in species with clumped mating, such as parasitoid wasps like Nasonia vitripennis, where females adjust sex ratios toward females with increasing foundress numbers, matching LMC predictions and reducing brother competition.^[11] Similarly, pollinating fig wasps (Agaonidae) exhibit extreme female biases (often 1:10 to 1:50 male:female) in singly founded figs, with ratios shifting toward equality as more foundresses compete.^[10] Female-biased ratios also occur in mites with patch-restricted dispersal, confirming LMC's role across taxa.^[9] Extensions of LMC include multi-generational models for persistent subdivided populations, such as haystack structures, where cumulative relatedness over generations can intensify female biases beyond single-generation predictions. Additionally, sex ratio distortion by males, as seen in the paternal sex ratio (PSR) chromosome of Nasonia vitripennis, creates intragenomic conflict by inducing all-male broods in infected females, countering LMC's female bias and reducing host fitness.^[12]

Local Resource Competition

In philopatric species, such as many birds and mammals, daughters tend to remain in or near their natal area, leading to intense competition with mothers and sisters for limited local resources like breeding territories or food, whereas sons often disperse and thus avoid such rivalry.^[13] This asymmetric competition favors the evolution of male-biased sex ratios, as producing more sons reduces future resource conflicts among female kin and thereby enhances the mother's inclusive fitness within the kin selection framework.^[13] The theoretical foundation for local resource competition (LRC) builds on an extension of kin selection models, as developed by Bull in 1983, where parents adjust offspring sex ratios to minimize the fitness costs imposed by resource scarcity on philopatric offspring. In this model, male-biased allocation evolves because the reproductive value of daughters is depressed by competitive interactions with relatives, prompting a shift away from the equal investment predicted by Fisher's principle under conditions of high female philopatry. LRC predicts stronger male biases in species exhibiting high female natal philopatry combined with resource limitation, as these factors amplify the fitness penalty for producing daughters.^[14] Qualitatively, the optimal sex ratio shifts toward males when competition suppresses female reproductive value, effectively lowering the relative benefits of investing in daughters compared to sons who disperse and face less kin-mediated rivalry. Empirical support for LRC comes from studies on red deer (Cervus elaphus), where high population density intensifies resource competition among philopatric females, correlating with male-biased offspring sex ratios that alleviate future maternal-daughter conflicts. Similarly, in Seychelles warblers (Acrocephalus sechellensis) under high-density conditions on poor-quality territories, parents produce more male offspring, consistent with LRC as females compete locally for limited breeding sites while males disperse more readily.^[15] The strength of the male bias under LRC diminishes when male philopatry increases, as sons then also contribute to local resource competition, reducing the relative advantage of producing them over daughters.^[14]

Local Resource Enhancement

Local resource enhancement (LRE) refers to the process in which siblings of one sex provide mutual benefits to the family group through cooperative behaviors, such as resource sharing, foraging assistance, or territory maintenance, thereby increasing the inclusive fitness returns from producing that sex. In species where females exhibit greater philopatry and contribute more substantially to these group-level benefits—often in cooperative breeding systems—parents gain higher fitness by allocating more resources to daughters, as their presence amplifies the productivity of the natal group for all relatives. This mechanism elevates the relative value of female offspring compared to males, who may disperse and contribute less to local cooperation.^[16] The theoretical foundation of LRE builds on extensions of kin selection theory, where female-biased philopatry fosters mutualistic interactions among relatives, enhancing overall group fitness beyond individual parental investment. In cooperative breeders with sex-specific roles, the inclusive fitness benefits from female helpers—such as improved sibling survival through shared provisioning—outweigh the costs of producing them, leading to evolutionary pressure for female-biased sex ratios. This contrasts with scenarios dominated by rivalry, where positive cooperative interactions exceed competitive costs, shifting allocation toward the sex that maximizes group enhancement. Seminal models incorporate these trade-offs qualitatively, showing that when the fitness increment from female cooperation surpasses that from male dispersal, optimal allocation favors daughters.^[16]^[17] Predictions from LRE include female-biased offspring sex ratios in species characterized by female-specific cooperative roles, limited female dispersal, and high potential for group resource augmentation, particularly in high-quality habitats where helpers can effectively contribute. For instance, in the Seychelles warbler (Acrocephalus sechellensis), a cooperatively breeding bird, parents on high-quality territories produce significantly more daughters (up to 80% female-biased broods), as retained female offspring assist in foraging and chick feeding, boosting subsequent reproductive success by approximately 90% (from 0.85 to 1.62 yearlings fledged) compared to groups without female helpers.^[15]^[18] However, a 2024 comparative analysis suggests that in the Seychelles warbler, the female bias may be more attributable to avoiding local resource competition than to enhancement from helping.^[19] Similarly, in the primitively social bee Exoneura bicolor, female-biased sex ratios (around 70% females) arise because coexisting sisters share nest resources and defend against intruders, enhancing colony productivity and inclusive fitness gains for the mother. These patterns hold when cooperation dominates over rivalry, as verified in comparative analyses across vertebrates and invertebrates.^[20]

Condition-Dependent Strategies

Trivers-Willard Hypothesis

The Trivers-Willard hypothesis, proposed by Robert Trivers and Dan Willard in 1973, posits that parents in good physiological condition should bias the production of offspring toward the sex exhibiting greater variance in reproductive success, typically males, while those in poor condition should favor the sex with more consistent returns, usually females.^[21] This condition-dependent strategy builds briefly on Fisher's principle of equal overall investment in both sexes across a population, but allows for facultative deviations when parental condition varies.^[21] The underlying logic stems from differences in how additional parental resources translate into offspring reproductive success across sexes. In many species, males benefit disproportionately from extra investment, such as larger body size leading to higher mating success through competition, whereas females often show more linear returns regardless of size due to less intense sexual selection.^[21] Thus, high-condition parents maximize fitness by allocating resources to sons, who can capitalize on the boost, while low-condition parents do better with daughters, whose success is less sensitive to reduced investment.^[21] Key assumptions include a positive correlation between parental condition and offspring viability or condition, and the presence of sexual selection that amplifies benefits for high-quality individuals of the variance-prone sex.^[21] The hypothesis further assumes that parents can adjust offspring sex ratios or investment in response to their condition, often in contexts of polygynous mating where male reproductive success varies widely.^[21] The mathematical framework conceptualizes optimal sex allocation as the strategy that maximizes parental expected fitness, given condition-dependent returns. If parental investment I influences male fitness more steeply than female fitness—e.g., male reproductive success R_m(I) rises nonlinearly while female R_f(I) is more constant—then for high I, the proportion allocated to sons p_m should exceed 0.5 to equate marginal fitness gains per unit investment across sexes.^[21] This ensures that the total fitness W = p_m R_m(I_m) + (1 - p_m) R_f(I_f), with I_m + I_f = I, is optimized.^[21] Foundational empirical support comes from studies on ungulates, such as red deer (Cervus elaphus), where dominant, high-ranking mothers produce more sons, and these sons achieve greater lifetime reproductive success due to enhanced competitive ability.^[22] Similar patterns appear in other polygynous mammals, like feral horses and bighorn sheep, linking maternal dominance or body condition to male-biased offspring ratios.^[21] Criticisms highlight that the hypothesis applies most robustly to polygynous species with high male variance in mating success, showing weaker or reversed effects in monogamous systems where sex differences in returns are minimal.^[23] Refinements emphasize distinguishing sex ratio biases at birth from post-birth investment adjustments, with rigorous tests requiring longitudinal data tracking parental condition, offspring sex, and lifetime reproductive outcomes to confirm fitness benefits.^[23]

Effects of Parental Condition

Parental condition, encompassing factors such as nutritional status and physiological health, influences sex allocation strategies in various species, often aligning with predictions from the Trivers-Willard hypothesis where parents in better condition bias investment toward the sex with greater reproductive potential.^[24] In birds, food availability plays a key role in shaping offspring sex ratios, with well-nourished mothers typically producing more sons, who often require greater resources due to larger size and higher energy demands. For instance, in the monogamous European shag (Phalacrocorax aristotelis), experimental food supplementation during breeding led parents to produce a higher proportion of male offspring compared to controls, as males were heavier and larger at fledging.^[25] Similarly, in blue tits (Cyanistes caeruleus), high-quality mothers in resource-rich environments overproduce sons to capitalize on their potential for higher reproductive success.^[26] Conversely, under low-resource conditions, mothers favor daughters, whose reproduction is more reliable and less variance-prone.^[24] In mammals, maternal condition and quality similarly drive biases toward sons among healthier females, reflecting the higher variance in male reproductive success. Healthier mammalian mothers, often measured by body mass or overall vigor, produce more male offspring to exploit opportunities for polygynous mating.^[27] Historical data from human populations, such as 18th- and 19th-century European records, show that women in good nutritional and socioeconomic condition bore more sons, while those in poor condition had more daughters.^[28] In nonhuman primates like rhesus macaques (Macaca mulatta), dominant or well-conditioned females exhibit a slight bias toward male offspring, though evidence is inconsistent across species due to varying social structures.^[29] Experimental manipulations confirm these patterns by altering resource availability or physiological state. In Japanese quail (Coturnix japonica), food restriction or elevated corticosterone levels (a stress hormone indicating poor condition) resulted in female-biased offspring sex ratios, while supplementation or reduced stress shifted toward males.^[30] In mice (Mus musculus), diets high in carbohydrates or calcium led to male-biased litters, whereas low-calcium or restricted feeding produced more females, demonstrating direct links between maternal nutrition and sex allocation.^[31]^[32] Condition is often quantified via body mass, with heavier mothers biasing toward sons, or hormones like testosterone, where elevated maternal levels around conception correlate with more male offspring in both birds and mammals.^[33]^[34] Recent studies highlight how climate-driven resource variation, such as droughts, exacerbates condition-dependent biases in wild populations. A 2023 analysis of painted turtles (Chrysemys picta) in long-term field data revealed that extreme drought conditions, reducing resource availability, led to female-biased hatchling sex ratios, potentially delaying female maturity but enhancing immediate survival.^[35] In mammals, a 2024 review of wild populations found limited but emerging evidence that drought-induced resource scarcity prompts female-biased allocation to mitigate risks in male-competitive environments.^[36] These condition-dependent strategies are not universal, particularly failing in monogamous species where low variance in male reproductive success reduces the adaptive value of biasing toward sons. In such systems, like many socially monogamous birds and contemporary human societies, sex ratios remain closer to 1:1 regardless of parental condition.^[37]

Mate Quality and Offspring Value

In sex allocation strategies, females often bias offspring sex ratios toward the sex that can best capitalize on the genetic benefits provided by high-quality mates, particularly when sons stand to inherit mating advantages from attractive or dominant sires. This adjustment aligns with condition-dependent frameworks where parental investment favors offspring with higher reproductive value. For instance, in species with strong sexual selection on male traits, producing more sons when paired with superior males maximizes indirect fitness gains through grandsons' success. Empirical evidence from birds illustrates this pattern clearly. In blue tits (Cyanistes caeruleus), females paired with males exhibiting brighter ultraviolet crown plumage—a key attractiveness signal—produce broods with a higher proportion of sons, as confirmed by molecular sexing techniques. Similarly, in peafowl (Pavo cristatus), females mated to males with more elaborate eye-spotted trains, indicative of superior genetic quality, lay eggs that develop into male-biased offspring, with yolk carotenoid levels correlating to this shift. These biases extend to dominance hierarchies; in red deer (Cervus elaphus), high-ranking males sire more sons, as daughters gain fewer benefits from inheriting competitive traits suited to male contests.^[38]^[39]^[40] Genetic paternity analyses in socially monogamous species further validate these mate-driven biases, distinguishing them from extra-pair influences. In collared flycatchers (Ficedula albicollis), microsatellite markers reveal that sex ratio adjustments occur primarily in within-pair offspring, with females producing more sons sired by socially attractive mates displaying large white wing patches, independent of genetic paternity deviations. This confirms behavioral assessment of mate quality via visual ornaments, rather than undetected cuckoldry. Trade-offs arise as females balance these adjustments against reproductive costs; for example, in blue tits, perceived mate quality via behavioral displays prompts son bias only when environmental cues signal low risk to male offspring survival, avoiding overinvestment in vulnerable sexes.^[41]^[42] Recent studies highlight molecular mechanisms linking mate compatibility to allocation. In three-spined sticklebacks (Gasterosteus aculeatus), a 2023 investigation found that females adjust sex ratios toward daughters when paired with MHC-dissimilar males, enhancing offspring immune diversity in parasite-rich environments, as determined by genotyping and controlled pairings. This demonstrates how females perceive mate quality through olfactory MHC cues, trading off sex bias for immunological benefits in aquatic systems.^[43]

Facultative Sex Change

Facultative sex change, also known as sequential hermaphroditism, enables organisms to alter their reproductive role from one sex to the other during their lifetime, optimizing fitness based on changing conditions such as body size or social status.^[44] In protandrous species, individuals begin life as males and transition to females, while in protogynous species, the reverse occurs, with females changing to males. This strategy is particularly prevalent in fish, where reproductive success often correlates with size; smaller individuals may benefit more from male roles due to lower costs of sperm production, whereas larger sizes enhance female fecundity through increased egg output.^[45] The timing of sex change aligns with the Trivers-Willard hypothesis, which posits that parents or individuals should bias allocation toward the sex offering higher fitness returns given current condition.^[46] In sequential hermaphrodites, this manifests as switching to the sex with greater reproductive value at a particular life stage; for instance, in many fish, juveniles start as the sex benefiting from early reproduction (often male), then shift to female as size increases and egg production becomes more advantageous.^[21] This condition-dependent adjustment maximizes lifetime reproductive success by exploiting size-related asymmetries in mating opportunities and parental investment.^[46] Theoretical models, such as those developed by Charnov and colleagues, predict that the optimal timing for sex change occurs when the marginal fitness gain from adopting the new sex exceeds the loss from ceasing reproduction in the current sex.^[47] These models emphasize a size-advantage framework, where the evolutionarily stable strategy involves changing sex at the point where lifetime fitness curves for male and female functions intersect, often favoring protogyny in species with male-biased operational sex ratios.^[48] Empirical support comes from coral reef fishes, where larger body size disproportionately boosts female reproductive output; for example, in protogynous species like groupers, mature females below size limits contribute significantly to population egg production, underscoring the selective pressure for delayed sex change until larger sizes.^[49]^[50] Prominent examples include the clownfish (Amphiprion spp.), which exhibit protandry: all individuals are born male, with the dominant individual developing into the sole female in a social group; if the female dies, the largest male rapidly transitions to female to maintain breeding.^[51] This change is socially triggered, as demonstrated in laboratory experiments where removal of the dominant female induces the sex reversal within weeks.^[52] Conversely, many wrasses, such as the cleaner wrasse (Labroides dimidiatus) and bluehead wrasse (Thalassoma bifasciatum), are protogynous; the largest female changes to male upon the dominant male's removal, with lab studies showing complete gonadal transformation in as little as 10 days under manipulated social conditions.^[53]^[54]^[55] Recent genomic studies (2022–2025) have advanced understanding of sex change triggers in aquaculture species, identifying key regulatory pathways responsive to environmental cues like temperature and social density. For instance, transcriptomic analyses in largefin longbarbel catfish (Clarias macrocephalus) reveal gene expression shifts during induced reversals, highlighting potential for selective breeding to control sex allocation in farming.^[56] Similarly, research on temperature-induced early sex change in ricefield eels (Monopterus albus) demonstrates skewed sex ratios under elevated temperatures, informing sustainable aquaculture practices.^[57]

Sex Allocation in Plants

Models for Hermaphroditic Plants

In hermaphroditic plants, which simultaneously perform male and female reproductive functions, sex allocation theory addresses how resources are optimally divided between pollen production (male function) and ovule or seed development (female function) to maximize overall fitness. This framework extends Fisher's principle of equal parental investment in separate-sexed offspring to cosexual organisms by treating male and female gamete production as analogous but concurrent contributions to fitness gains through siring and maternity.^[58] Pioneered by Eric L. Charnov in his 1982 monograph, these models emphasize that plants evolve to balance allocation such that incremental investments in each function yield equivalent fitness returns, accounting for the unique constraints of pollen dispersal and local mating dynamics in plants.^[58] The core prediction derives from an evolutionarily stable strategy (ESS) approach, where optimal resource allocation R_m to male function and R_f to female function equalizes the marginal fitness gains per unit resource. This occurs when the rate of change in male fitness F_m with respect to male allocation equals that of female fitness F_f with respect to female allocation:

\frac{dF_m}{dR_m} = \frac{dF_f}{dR_f}

Such equilibrium often results in equal total allocation to both functions under symmetric returns, but asymmetries in pollen transfer efficiency or selfing rates shift the balance.^[58] Key factors influencing these returns include pollen discounting, where increased self-pollen production on stigmas reduces the relative success of exported pollen, thereby diminishing marginal gains from male investment, and geitonogamy, the within-plant pollen transfer that exacerbates selfing and further discounts male fitness in large inflorescences.^[59] These models yield specific predictions for allocation patterns: outcrossing hermaphrodites tend toward male-biased allocation due to higher returns from pollen dispersal in open populations, while selfing species favor female bias as male function yields diminishing benefits from local mating.^[58] Temporal adjustments, such as protandry (male function preceding female in flowers), further optimize allocation by minimizing geitonogamy and enhancing outcrossed siring early in anthesis.^[60] For instance, in sequentially blooming cosexual plants like those in the genus Ipomopsis, flowers adjust allocation based on pollinator visitation patterns; early-blooming flowers invest more in pollen presentation when pollinators carry less foreign pollen, while later flowers shift toward ovule production as visitation efficiency for male function declines.^[60] Recent studies have addressed gaps in these models regarding rapid evolutionary shifts post-genomic events, demonstrating that whole-genome duplication (WGD) can quickly favor male-biased allocation in neo-polyploids. In the hermaphroditic perennial Galax urceolata, neo-autotetraploid populations evolved significantly higher male allocation compared to diploid progenitors within a few generations, likely due to altered dosage effects on reproductive traits that accelerate male fitness gains under the ESS framework.^[61]

Environmental Influences on Plant Sex Expression

In dioecious plants, environmental sex determination (ESD) allows external factors such as temperature, light, and nutrient availability to influence the expression of male or female phenotypes, often during critical developmental stages. For instance, elevated temperatures can bias sex ratios toward males in certain tree species, as males typically exhibit greater tolerance to heat stress due to differences in resource acquisition and physiological responses. ^[62] Similarly, light intensity affects sex expression in Equisetum (horsetails), where high light levels promote female gametophyte development, while low light favors males, reflecting adaptive strategies to optimize reproduction under varying irradiance. ^[63] Resource-dependent biases, such as those under nutrient scarcity, can lead to female-biased sex ratios, aligning with theoretical predictions from sex allocation models that emphasize condition-dependent strategies. ^[64] Empirical studies highlight dynamic environmental influences on sex expression, including seasonal shifts and responses to global change. In striped maple (Acer pensylvanicum), a polygamo-dioecious species, trees frequently change sex expression annually, with healthier individuals more likely to express as males in favorable seasons, while stressed trees shift toward female or non-reproductive states, potentially as a bet-hedging strategy against mortality. ^[65] Recent research on dioecious plants indicates that climate-induced stressors, such as increased aridity and warming, can exacerbate male biases in sex ratios by differentially affecting female survival and reproduction, leading to potential population declines in altered habitats. ^[66] These patterns underscore how abiotic factors interact with plant physiology to modulate sex allocation, often without altering underlying genetic sex determination. In monoecious plants, environmental cues enable flexible adjustment of male-to-female flower ratios on the same individual, enhancing reproductive efficiency. For example, in maize (Zea mays), nutrient availability and photoperiod influence tassel (male inflorescence) size and silk (female) development, with high resource levels promoting larger male structures to maximize pollen dispersal under optimal conditions. ^[67] Pollinator availability further interacts with these adjustments in hermaphroditic or monoecious systems, where reduced pollinator density prompts increased allocation to female function to capitalize on selfing or limited mating opportunities, as observed in insect-pollinated perennials. ^[68] Ongoing research identifies critical gaps in understanding how global change disrupts plant sex ratios, particularly through compounded effects of drought, elevated CO2, and temperature. A 2025 review emphasizes that anthropogenic stressors aggravate water limitations, altering sex expression in dioecious and monoecious species and potentially leading to skewed ratios that impair long-term population viability. ^[69] These disruptions highlight the need for integrated studies on environmental thresholds to predict resilience in plant reproductive strategies under future climates.

Proximate Mechanisms

Genetic and Genomic Controls

Sex determination systems form the genetic foundation of sex allocation across taxa, influencing primary sex ratios through chromosomal mechanisms. In many animals, the XY system predominates, where males are heterogametic (XY) and the Y chromosome typically carries male-determining genes, such as SRY in mammals.^[70] Conversely, birds and some reptiles employ the ZW system, with females heterogametic (ZW) and males homogametic (ZZ); dosage compensation on the Z chromosome helps regulate sex-specific gene expression, potentially modulating allocation biases.^[70] Plants exhibit diverse systems, including XY-like arrangements in dioecious species such as papaya, where sex chromosomes evolve from autosomes via mutations causing sterility in one sex.^[70] These chromosomal setups constrain or enable sex ratio variation, with heterogamety allowing for potential biases in gamete production or viability. Haplodiploidy in insects, particularly Hymenoptera (e.g., bees and wasps), provides a unique genetic basis for sex allocation biases under local mate competition (LMC). In this system, males develop from unfertilized eggs (haploid) and females from fertilized eggs (diploid), enabling mothers to adjust sex ratios by controlling sperm release; this favors female-biased allocation in structured populations where brothers compete locally, aligning with Hamilton's LMC theory. Such genetic control amplifies relatedness asymmetries, promoting eusociality and precise sex ratio adjustment to optimize inclusive fitness. Genomic imprinting introduces parent-of-origin effects that fuel intragenomic conflicts over sex allocation, particularly in mammals. Paternally expressed imprinted genes often promote greater resource allocation to male offspring, enhancing paternal fitness through increased male reproductive success, as seen in models where such genes bias embryonic growth or behavior toward sons in polygynous systems. This conflict arises because paternally inherited alleles favor investment in larger, more competitive sons, while maternally inherited alleles prioritize even allocation to avoid exploitation; empirical evidence from imprinted loci like Igf2 supports this manipulation in rodent models. Quantitative trait locus (QTL) mapping and genome-wide association studies (GWAS) have identified genetic architectures underlying sex allocation biases. In Drosophila simulans, QTL analysis of recombinant inbred lines revealed five major loci across chromosomes 2 and 3 that suppress X-linked meiotic drive, restoring balanced sex ratios by countering female-biased distortion; these loci explain over 50% of phenotypic variance through epistatic interactions.^[71] Meiotic drive elements further distort allocation by biasing sex chromosome transmission, such as X-linked drivers in Drosophila that eliminate Y-bearing sperm, leading to female-biased ratios unless suppressed by autosomal modifiers.^[72] Recent genomic advances highlight dynamic evolutionary shifts in sex allocation. A 2025 study on the hermaphroditic plant Galax urceolata demonstrated that whole-genome duplication in neo-autotetraploids facilitates rapid evolution of male-biased allocation, with duplicated genomes allowing relaxed trade-offs between male and female function compared to diploid progenitors.^[61] This polyploidy enables higher male gamete investment without compromising female fertility, accelerating adaptation in changing environments.^[61] Genetic variation in sex allocation loci maintains evolutionary plasticity, permitting populations to respond to selective pressures like resource availability or mating systems. Such standing variation, often polygenic, buffers against fixation of extreme biases and supports condition-dependent strategies, as modeled in systems where heritable plasticity enhances long-term fitness under variable environments.^[73] This genetic underpinnings ensures sex allocation evolves as a flexible trait, balancing conflicts and promoting adaptive divergence across taxa.^[73]

Environmental and Physiological Adjustments

Organisms adjust sex allocation through environmental and physiological mechanisms that enable rapid, reversible responses to external cues, distinct from fixed genetic programs. Key cues include hormonal signals, such as varying estrogen-to-testosterone ratios in vertebrates, which influence gonadal development and offspring sex ratios during critical reproductive windows.^[74] Photoperiod, or day length, serves as another cue, particularly in fish and invertebrates, where longer days promote female-biased allocation by modulating reproductive timing and hormone release.^[75] Population density also acts as a cue, triggering density-dependent adjustments where higher densities often lead to female-biased sex ratios to mitigate intense local competition among males.^[76] Physiological pathways translate these cues into sex allocation changes without altering the genome. In reptiles exhibiting temperature-dependent sex determination, nest temperature is sensed via calcium influx in embryonic gonadal cells, activating temperature-sensitive proteins like phosphorylated STAT3, which represses genes favoring male development at warmer temperatures (around 31°C), thereby promoting ovarian differentiation.^[77] Behavioral adjustments further facilitate these responses; for instance, in insects under local mate competition, females detect crowding through host patch cues or prior oviposition signals and bias clutches toward daughters by adjusting egg fertilization timing or sequence, reducing male-male rivalry in dense patches.^[78] Specific examples illustrate this plasticity: in turtles like the red-eared slider (Trachemys scripta elegans), nest temperatures below 26°C produce all-male clutches, while those above 31°C yield all-female offspring, shifting population sex ratios based on incubation conditions.^[79] Similarly, in nematodes such as Caenorhabditis species, pheromone detection signals population density or mate availability, modulating dauer formation and subsequent sex allocation to favor reproductive output under stress.^[80] The evolution of such facultative plasticity balances costs and benefits tied to environmental variability. Benefits include enhanced fitness in fluctuating conditions, as seen in hermaphroditic flatworms where individuals increase male gamete allocation in response to higher mating group sizes, optimizing fertilization success amid variable sperm competition.^[81] However, costs arise from resource trade-offs, such as reduced overall fecundity or survival when plasticity mechanisms demand energy for cue sensing and rapid physiological shifts, particularly in stable environments where fixed strategies suffice.^[81] Recent research highlights epigenetic modifications as bridges linking environmental cues to sex allocation, especially under climate change pressures. In various animals, including reptiles and fish, temperature-induced epigenetic changes, like DNA methylation patterns in sex-determining genes, alter allocation across generations, with warmer conditions often feminizing populations and exacerbating imbalances from rising global temperatures.^[82] Multigenerational studies show that paternal exposure to elevated temperatures modifies offspring epigenetic profiles, influencing sex development pathways and potentially amplifying climate-driven shifts in allocation.^[83]

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