Age-Related Loss of Lumbar Spinal Lordosis and Mobility – A Study of 323 Asymptomatic Volunteers

Background

The understanding of the individual shape and mobility of the lumbar spine are key factors for the prevention and treatment of low back pain. The influence of age and sex on the total lumbar lordosis and the range of motion as well as on different lumbar sub-regions (lower, middle and upper lordosis) in asymptomatic subjects still merits discussion, since it is essential for patient-specific treatment and evidence-based distinction between painful degenerative pathologies and asymptomatic aging.

Methods and Findings

A novel non-invasive measuring system was used to assess the total and local lumbar shape and its mobility of 323 asymptomatic volunteers (age: 20–75 yrs; BMI <26.0 kg/m²; males/females: 139/184). The lumbar lordosis for standing and the range of motion for maximal upper body flexion (RoF) and extension (RoE) were determined. The total lordosis was significantly reduced by approximately 20%, the RoF by 12% and the RoE by 31% in the oldest (>50 yrs) compared to the youngest age cohort (20–29 yrs). Locally, these decreases mostly occurred in the middle part of the lordosis and less towards the lumbo-sacral and thoraco-lumbar transitions. The sex only affected the RoE.

Conclusions

During aging, the lower lumbar spine retains its lordosis and mobility, whereas the middle part flattens and becomes less mobile. These findings lay the ground for a better understanding of the incidence of level- and age-dependent spinal disorders, and may have important implications for the clinical long-term success of different surgical interventions.     

Blood pressure levels of the Suruí and Zoró Indians of the Brazilian Amazon: group- and sex-specific effects resulting from body composition, health status, and age

Two related Tupí-Mondê-speaking tribes of the Aripuan? Indian Park of western Brazil are compared in terms of their recent contact with Western culture, subsistence patterns, general health, and blood pressure levels. Age, weight, height, sex, and tribal affiliation for Suruí and Zoró adults over age 18 are included in an analysis of covariance to test regression models of both diastolic and systolic blood pressure. Because of significant interaction effects between sex and other covariates, sex-specific models were developed. The relationship between body mass and blood pressure level in males conforms with Western data, but the direction and magnitude of effects for the age and body mass covariates in both sexes conflict with data from other modernizing societies. With age, weight/height ratio, and sex differences controlled for, Suruí males show a lower mean systolic blood pressure (SBP) level and diastolic blood pressure (DBP) level than Zoró males. Intertribal differences were smaller among females: Suruí females SBP and DBP differences were similar but did not reach significance. Other sex-specific differences include a greater inverse relationship between age and SBP level among the 104 male subjects than among the 98 female subjects (with similar trends in DBP) and a small but significant effect of the weight/height ratio on both SBP and DBP in males but not in females. Health status data for these groups suggest that hypothesized increases in mean blood pressure levels following the Suruí's acceptance of a Western diet and social stratification may be modified by their health status, particularly prevalent infectious disease.     

Day-old chicks categorised on latency to peck, exhibit a stable fear pattern until 15 days of age

Day-old chicks of both sexes were individually categorised on the latency to peck pebbles and termed as high latency (HL), moderate latency (ML) or low latency (LL). Anxiolytic doses of diazepam diminished the latency only in the HL category, suggesting that it is the most anxious category. At 15 days of age, the LL category showed the lowest latency to ambulate in the open-field test, the lowest immobility duration in the tonic immobility test and insensitivity to anxiolytic doses of diazepam in both behavioural tests suggesting that it is the less anxious category. The increase of the central benzodiazepine receptor density induced by acute stressors was the highest in the most anxious and/or fearful HL category. There were more females than males in the LL category and inversely in the HL category there were more males. The results suggest that the fear pattern depends on the sex and inter-individual differences within a same sex which are stable across life. This could be used as a test for fear and/or anxiety state, and useful to choose fowls with the best performance later in life.     

Latitudinal variation in sexual dimorphism in life‐history traits of a freshwater fish

Sexual dimorphism is common across the animal kingdom, but the contribution of environmental factors shaping differences between the sexes remains controversial. In ectotherms, life‐history traits are known to correlate with latitude, but sex‐specific responses are not well understood. We analyzed life‐history trait variation between the sexes of European perch (Perca fluviatilis L.), a common freshwater fish displaying larger female size, by employing a wide latitudinal gradient. We expected to find sex‐dependent latitudinal variation in life‐history variables: length at age, length increment, and size at maturity, with females showing consistently higher values than males at all latitudes. We further anticipated that this gender difference would progressively decrease with the increasingly harsh environmental conditions toward higher latitude. We hypothesized that growth and length increment would decrease and size/age at maturity would increase at higher latitudes. Our results confirmed female‐biased sexual size dimorphism at all latitudes and the magnitude of sexual dimorphism diminished with increase in latitude. Growth of both sexes decreased with increase in latitude, and the female latitudinal clines were steeper than those of males. Hence, we challenge two predominant ecological rules (Rensch's and Bergmann's rules) that describe common large‐scale patterns of body size variation. Our data demonstrate that these two rules are not universally applicable in ectotherms or female‐biased species. Our study highlights the importance of sex‐specific differences in life‐history traits along a latitudinal gradient, with evident implications for a wide range of studies from individual to ecosystems level.     

Anthropometric variationin the Yanadi: Population structure and sex differences

Anthropometric variation and sex differences were investigated among the Yanadi tribe, who live in different regions and show differences in population structure variables and form regional breeding populations. In case of within population variation in anthropometric characters, both males and females show greater variation in a few specific characters (e.g., HB, LL) and almost no variation in FB, but least variation in HL and NB in males and in ZB, LB in females and show sex differences. Overall, females showed greater variation in more number of characters than males. In case of between population variation a few traits show clinal trend between male-female comparisons. Each character shows a specific pattern (e.g., HB, ZB, EL, GB) which vary according to the spatial distribution of the regional populations. The curves also indicate least average differences corresponding to within regional homogeneity among males and females (e.g., HB, ECD, EB) and increasing differences with males and females of other regions. A comparison of anthropometric profiles of the five populations show significant sex differences in IY subpopulation. The three mainland subpopulations show wide morphological differentiation with two coastal subpopulations, who also differ in subsistence pattern and geographically isolated. A comparison of anthropometric profiles between males and females across five populations show positive association between P1, HF and P₂ populations, whereas significant negative association with CY and HF. The correllelographs based on the male-female comparisons also indicate greater morphological similarity between sexes in cases of within regional population. They also show clinal changes of either decreasing or increasing trends of morphological variation between populations in association with spatial distribution and population structure differences. The results obtained are in agreement with the expectations from the population structure of the tribe.     

The heritability of insomnia: A meta-analysis of twin studies

Twin studies of insomnia exhibit heterogeneity in estimates of heritability. This heterogeneity is likely because of sex differences, age of the sample, the reporter and the definition of insomnia. The aim of the present study was to systematically search the literature for twin studies investigating insomnia disorder and insomnia symptoms and to meta-analyse the estimates of heritability derived from these studies to generate an overall estimate of heritability. We further examined whether heritability was moderated by sex, age, reporter and insomnia symptom. A systematic literature search of five online databases was completed on 24 January 2020. Two authors independently screened 5644 abstracts, and 160 complete papers for the inclusion criteria of twin studies from the general population reporting heritability statistics on insomnia or insomnia symptoms, written in English, reporting data from independent studies. We ultimately included 12 papers in the meta-analysis. The meta-analysis focussed on twin intra-class correlations for monozygotic and dizygotic twins. Based on these intra-class correlations, the meta-analytic estimate of heritability was estimated at 40%. Moderator analyses showed stronger heritability in females than males; and for parent-reported insomnia symptoms compared with self-reported insomnia symptoms. There were no other significant moderator effects, although this is likely because of the small number of studies that were comparable across levels of the moderators. Our meta-analysis provides a robust estimate of the heritability of insomnia, which can inform future research aiming to uncover molecular genetic factors involved in insomnia vulnerability.     

Influence of age,sex, and caging on joint mobility in the patas monkey (Erythrocebus patas)

Normal range of joint mobility in the extremities of the patas monkey, Erythrocebus patas, was established for a free-ranging colony of 64 animals at La Parguera, Puerto Rico (Caribbean Primate Research Center). Eighty-five animals that had been caged (30″ × 30″ cages) for up to 5 years were used for comparison. Passive joint mobility of anesthetized animals was measured with a goniometer. Nine parameters (five on the forelimb and four on the hindlimb) were measured on each animal. The data were sorted into subsets according to the animal's age, sex, place of birth, and type of confinement, if any. The number of animals in each subset was recorded and the mean (in degrees) and standard deviation for each parameter were calculated. A P?0.05 on two-tailed Student's t-tests was considered significant. Comparisons between free-ranging males and females showed significant differences in one or two parameters for all age groups. A cross-sectional sample of free-ranging animals of both sexes showed that significant changes in joint mobility occurred only in the first 18 months of life. Joint mobility of all caged animals, however, was highly variable, and even between the more mature animals there were significant differences in several parameters. Almost all comparisons of subsets of the same age and sex showed significant differences between caged and free-ranging animals in at least one parameter. When the caged animals were laboratory-born, however, these differences were significant in five out of nine parameters. The results suggest that, although caging itself affects joint mobility, the age of first confinement may have an even greater effect than the length of the confinement.     

Universal sex differences in online advertisers age preferences: comparing data from 14 cultures and 2 religious groups

The search for a potential partner has been aided over recent years by the widespread use of online dating sites and this process of relationship formation has conveniently presented an ideal opportunity for researchers to analyze human mating desires and to compare evolutionary and social constructivist based hypotheses. One such aspect of human mating behaviour yet to be thoroughly explored using access to online dating advertisements is the idealized age desired by each sex when considering a possible relationship. This study accessed minimum (youngest age considered) and maximum (oldest age considered) age preferences from 14 separate cultures and two religious groups from both sexes at ages 20, 25, 30, 35, 40, 45, and 50 years. The results showed that overall there was a growing disparity between males own age and preferred age of partner as males themselves aged (as indicated by greater effect sizes with advertisers age), with females showing a pattern for preferences around their own age or older. Females did not express an age preference for males younger than male's age preferences for females at any advertiser's age. On only three occasions were there no age differences between the sexes in their desire to initiate a relationship with the opposite sex. The results were clearly concurrent with earlier findings supportive of evolutionary or adaptationist interpretations. Neither a random pattern of age preferences more consistent with an arbitrary norms prediction, nor clear evidence for toy boy proclivities in females or males was found. Future studies utilizing the methodology used in this study to examine other human mating decision making processes are proposed.     

Patterns of femoral bone remodeling dynamics in a medieval Nubian population

The relationship between age, sex and histomorphometry in femoral cortical bone was examined in a skeletal population of late Medieval antiquity (AD 1250–1450) from Kulubnarti, in Sudanese Nubia. These skeletal remains are naturally mummified and in an excellent state of preservation. The study sample consisted of femoral cross sections from 24 females and 19 males ranging in age from 20 to 50+ years. Femoral cross sections were examined using an image analysis system. Numbers of secondary osteons and osteon fragments were counted, osteon area and Haversian canal area were measured, and several variables were calculated to assess differences between sexes and among age groups in bone remodeling variables. The results indicate significant differences between the sexes in osteon number and size. Males had significantly more intact osteons than females, whereas females had significantly larger osteons than males. Haversian canal dimensions were not statistically significant between the sexes. Sex differences in activity patterns in which males were involved in more physically strenuous tasks may have contributed to differences in remodeling variables. Interpopulational comparisons suggest that mechanical strain affects the microstructural features examined in this study. In particular, small Haversian canals in some archaeological skeletal populations are associated with higher bone volume, which may result from high levels of mechanical strain. Am J Phys Anthropol 104:133–146, 1997. © 1997 Wiley-Liss, Inc.     

No differential consequences of reproduction according to sex in Juniperus communis var. depressa (Cupressaceae)

In dioecious plant species, males and females are thought to have dissimilar allocation patterns. Females are believed to invest more in reproduction and less in growth and maintenance than males. This differential investment between sexes could result in distinct growth patterns and contrasting survival rates, thereby affecting the sex ratio of a population and the age and size distribution of males and females, possibly leading to habitat segregation according to sex. These effects might become more apparent under particularly limiting conditions, such as in nutrient-deficient soils or in climatically stressed environments. To verify these predictions, growth patterns, microsite characteristics, and age and size distribution of male and female individuals were compared, and population sex ratio was determined in three populations of the dioecious shrub Juniperus communis var. depressa (Cupressaceae, Pinophyta) along a short latitudinal gradient on the eastern coast of Hudson Bay (Northern Québec, Canada). We found that the northernmost population had a male-biased sex ratio, but that the southernmost one had a higher proportion of females. Our results failed to reveal any significant differences in radial growth patterns, mean sensitivity, annual elongation of the main axis, and size and age frequency distribution between males and females in any population. Furthermore, there was no evidence of microhabitat segregation according to sex as indicated by the lack of differences in the physicochemical characteristics of the substrate under males and females. Clearly, the expected ecological consequences of a presumed greater investment of females in reproduction were not apparent even under the very stressful conditions prevailing on subarctic dunes. Many factors could reduce differences in the cost of reproduction between males and females, such as the number and quality of reproductive structures produced annually by individuals of each sex, the possible photosynthetic activity of the immature female cones, and the complexity of the source/sink relationship within individuals. Alternatively, there may be no differences between sexes in their reproductive investment.     

Body size dimorphism and sexual segregation in polygynous ungulates: an experimental test with Soay sheep

Sexual segregation in Soay sheep (Ovis aries) was investigated using an experimental approach in order to test the sexual dimorphism-body size hypothesis. Two corollaries of the sexual dimorphism-body size hypothesis were tested: (1) in dimorphic species males, the larger sex, have relatively smaller bite sizes on short swards because of the scaling of incisor arcade with body weight, and (2) they move off earlier to feed on taller but poorer-quality swards when such swards are patchily distributed on a scale which enables the spatial segregation of individuals. Patch choice between sexes was estimated using a matrix of grass patches which differed in both quality and biomass of grass on offer (HQ: high-quality-low-biomass; LQ: low-quality-high-biomass). Sex differences in patch choice and grazing behaviour were tested in short-term preference trials. Incisor breadth showed no significant difference between sexes. On the other hand, muzzle width was dimorphic, with females having a narrower muzzle than males. Bite size was significantly different between the sexes, being smaller in females than in males, although it was not significantly different between sward types. Females had a higher bite rate than males and the bite rate was higher in the HQ sward type than the LQ sward type. When the effect of body mass was removed, no sex differences in muzzle size, bite size or bite rate were found. The intake rate did not differ between the sexes or between sward types. Whilst both sexes preferred the HQ sward type, females spent a significantly longer time feeding on the LQ sward type than did males. The difference detected between the sexes in patch choice was not consistent directly with the sexual dimorphism-body size hypothesis. Alternative explanations based on sex differences in foraging behaviour in relation to body mass sexual dimorphism are discussed to explain the result. Received: 1 February 1999 / Accepted: 12 May 1999     

Sexing greater flamingos by weight and linear measurements

The sex of 59 adult Greater Flamingos (Phoenicopterus ruber roseus L.) was determined by laparoscopic exam of the gonads. Concomitant body weight (kg) and linear measurements (mm) of the culmen (bill), tarsus, middle toe, and wing were taken and compared for males vs. females. Although an overlap between sexes existed in all measurements, males on average were larger than females. Student's t-test indicated significant sexual differences for all five measurements. Thus, weights and linear measurements—especially tarsus, middle toe, and wing length—appear to be a useful parameter in determining an individual's sex in lieu of laparoscopic, feather pulp, blood chromosome, or fecal steroid analysis.     

Sex ratio and gender-dependent neighboring effects in <Emphasis Type="Italic">Podocarpus nagi</Emphasis>, a dioecious tree

We analyzed sex ratio, growth rates, and spacing among individuals of Podocarpus nagi, a dioecious tree, on Mt. Mikasa, Nara City, Japan. The sex ratio of reproductive trees ≥ 5 cm in stem diameter at breast height (dbh, 130 cm above ground level) was significantly male-biased. The sex ratio was male-biased in the < 20 cm and ≥ 50 cm size classes, while it did not depart from 1:1 in the 20 ≤ dbh < 50 cm class. Growth rate varied with tree size in males but not in females. The precocity and vigor of males suggests that differences in reproductive costs between sexes induce the biased sex ratio. Random labeling tests on the positions of reproductive trees showed that in the < 30 cm class, males and females were distributed randomly and independently from each other. In the ≥ 30 cm class, males were significantly clumped, whereas females were randomly distributed. Males and females showed significant repulsion, i.e., a spatial segregation of sexes. Both intra- and intersexual effects on the growth rate of crowding by neighbors were significant for females, but not for males. Maximum competitive interference was observed at a distance of 5 m, which corresponded approximately to the radius of clumps of large males and to the significant repulsive distance between large males and females. These results suggest that sexual differences in sensitivity to local crowding are related to the formation of gender-dependent spatial patterns. Formation of female-repulsive male clumps and a male-biased sex ratio may intensify the decreased probability of regeneration near males, as suggested by the limited seed-dispersal range of this species, thereby promoting coexistence with other species.     

Sex differences in telomeres and lifespan in Soay sheep: From the beginning to the end

There is tremendous diversity in ageing rates and lifespan not only among taxa but within species, and particularly between the sexes. Women often live longer than men, and considerable research on this topic has revealed some of the potential biological, psychological and cultural causes of sex differences in human ageing and lifespan. However, sex differences in lifespan are widespread in nonhuman animals suggesting biology plays a prominent role in variation in ageing and lifespan. Recently, evolutionary biologists have borrowed techniques from biomedicine to identify whether similar mechanisms causing or contributing to variation in ageing and lifespan in humans and laboratory animals also operate in wild animals. Telomeres are repetitive noncoding DNA sequences capping the ends of chromosomes that are important for chromosomal stability but that can shorten during normal cell division and exposure to stress. Telomere shortening is hypothesized to directly contribute to the ageing process as once telomeres shorten to some length, the cells stop dividing and die. Men tend to have shorter telomeres and faster rates of telomere attrition with age than women, suggesting one possible biological cause of sex differences in lifespan. In this issue of Molecular Ecology, Watson et al. ( 2017 ) show that telomere lengths in wild Soay sheep are similar between females and males near the beginning of life but quickly diverge with age because males but not females showed reduced telomere lengths at older ages. The authors further show that some of the observed sex difference in telomere lengths in old age may be due to male investment in horn growth earlier in life, suggesting that sexually dimorphic allocation to traits involved in sexual selection might underlie sex differences in telomere attrition. This study provides a rare example of how biological mechanisms potentially contributing to sex differences in lifespan in humans may also operate in free‐living animals. However, future studies using a longitudinal approach are necessary to confirm these observations and identify the ultimate and proximate causes of any sex differences in telomere lengths. Collaborations between evolutionary biologists and gerontologists are especially needed to identify whether telomere lengths have a causal role in ageing, particularly in natural conditions, and whether this directly contributes to sex differences in lifespan.     

The genetic architecture of behavioral traits in a spider

The existence of consistent individual differences in behavior has been shown in a number of species, and several studies have found observable sex differences in these behaviors, yet their evolutionary implications remain unclear. Understanding the evolutionary dynamics of behavioral traits requires knowledge of their genetic architectures and whether this architecture differs between the sexes. We conducted a quantitative genetic study in a sexually size‐dimorphic spider, Larinioides sclopetarius, which exhibits sex differences in adult lifestyles. We observed pedigreed spiders for aggression, activity, exploration, and boldness and used animal models to disentangle genetic and environmental influences on these behaviors. We detected trends toward (i) higher additive genetic variances in aggression, activity, and exploration in males than females, and (ii) difference in variances due to common environment/maternal effects, permanent environment and residual variance in aggression and activity with the first two variances being higher in males for both behaviors. We found no sex differences in the amount of genetic and environmental variance in boldness. The mean heritability estimates of aggression, activity, exploration, and boldness range from 0.039 to 0.222 with no sizeable differences between females and males. We note that the credible intervals of the estimates are large, implying a high degree of uncertainty, which disallow a robust conclusion of sex differences in the quantitative genetic estimates. However, the observed estimates suggest that sex differences in the quantitative genetic architecture of the behaviors cannot be ruled out. Notably, the present study suggests that genetic underpinnings of behaviors may differ between sexes and it thus underscores the importance of taking sex differences into account in quantitative genetic studies.     

Sex‐specific effects of protein and carbohydrate intake on reproduction but not lifespan in Drosophila melanogaster

Modest dietary restriction extends lifespan (LS) in a diverse range of taxa and typically has a larger effect in females than males. Traditionally, this has been attributed to a stronger trade‐off between LS and reproduction in females than in males that is mediated by the intake of calories. Recent studies, however, suggest that it is the intake of specific nutrients that extends LS and mediates this trade‐off. Here, we used the geometric framework (GF) to examine the sex‐specific effects of protein (P) and carbohydrate (C) intake on LS and reproduction in Drosophila melanogaster. We found that LS was maximized at a high intake of C and a low intake of P in both sexes, whereas nutrient intake had divergent effects on reproduction. Male offspring production rate and LS were maximized at the same intake of nutrients, whereas female egg production rate was maximized at a high intake of diets with a P:C ratio of 1:2. This resulted in larger differences in nutrient‐dependent optima for LS and reproduction in females than in males, as well as an optimal intake of nutrients for lifetime reproduction that differed between the sexes. Under dietary choice, the sexes followed similar feeding trajectories regulated around a P:C ratio of 1:4. Consequently, neither sex reached their nutritional optimum for lifetime reproduction, suggesting intralocus sexual conflict over nutrient optimization. Our study shows clear sex differences in the nutritional requirements of reproduction in D. melanogaster and joins the growing list of studies challenging the role of caloric restriction in extending LS.     

The influence of sex and body size on food habits of a giant tropical snake, Python reticulatus

1. In many animal species, dietary habits shift with body size, and differ between the sexes. However, the intraspecific range of body sizes is usually low, making it difficult to quantify size-associated trophic shifts, or to determine the degree to which sex differences in diet are due to body-size differences. Large snakes are ideal for such a study, because they provide a vast range of body sizes within a single population.
2. More than 1000 Reticulated Pythons ( Python reticulatus ) from southern Sumatra were examined, with specimens from 1·5 to > 6 m in snout–vent length, and from 1 to 75 kg in mass. Females attained much larger body sizes than did conspecific males (maxima of 20 vs 75 kg, 5 vs 7 m), but had similar head lengths at the same body lengths.
3. Prey sizes, feeding frequencies and numbers of stomach parasites (ascarid nematodes) increased with body size in both sexes, and dietary composition changed ontogenetically. Small snakes fed mostly on rats, but shifted to larger mammalian taxa (e.g. pangolins, porcupines, monkeys, wild pigs, mouse deer) at 3–4-m body length.
4. Adult males and females showed strong ecological divergence. For some traits, this divergence was entirely caused by the strong allometry (combined with sexual size dimorphism), but in other cases (e.g. feeding frequency, dietary composition), the sexes followed different allometric trajectories. For example, females shifted from rats to larger mammals at a smaller body size than did conspecific males, and feeding frequencies increased more rapidly with body size in females than in males. These allometric divergences enhanced the degree of sex difference in trophic ecology induced by sexual size dimorphism.     

ORIGINAL ARTICLE: Leptin associations with age,weight, and sex among chimpanzees (Pan troglodytes)

Background Leptin is a hormone secreted primarily by adipocytes, a lipostatic signal to the hypothalamus, and is often correlated with adiposity. Associations between leptin, age, and development are unknown in human’s closest evolutionary relative, the common chimpanzee (Pan troglodytes). Methods Serum leptin was assessed cross sectionally in association with age, weight, and sex in healthy captive chimpanzee males (n = 47) and females (n = 49) to test hypotheses related to predicted differences in leptin levels with body mass, development, and sexual dimorphism. Results Leptin increased with age and weight among females, but not in males. Leptin was overall higher in females compared to males. Conclusions Sex differences in leptin were most evident during adolescence and adulthood, despite similar increases in weight in both sexes indicating that sexual maturation is a key divergence point for differential somatic investment in adiposity and leptin levels between male and female chimpanzees.     

Asymmetrical expression of transsexual phenotypes in hummingbirds

I present evidence for asymmetry in the expression of transsexual traits in adult hummingbirds. Among females, individuals with male-like plumage are common and define a continuous range of variation. Among males, individuals with female-like plumage are rare and define discontinuous plumage morphs. Quantitative characters also distinguish transsexuals from other members of their sex, but the characters involved differ for male-like females (bill length) and female-like males (bill, wing and tail length). Gonadal development is correlated with transsexuality only in males; female-like males have significantly smaller testes than male-like males. Both sexes demonstrate a significant negative association between plumage brightness and bill length. This association suggests an ecological basis for transsexuality because differences in plumage and bill morphology are associated with differences in foraging behaviour within and between hummingbird species. Morphological differences between transsexuals and non-transsexuals imply that plumage sexual dimorphism is more likely to evolve through changes in the frequency of female, rather than male, transsexual variants.