Carotenoid status signaling in captive and wild red-collared widowbirds: independent effects of badge size and color

Carotenoid-based plumage ornaments are typically consideredto be sexually selected traits, functioning as honest condition-dependentsignals of phenotypic quality, but few studies have addressedthe function of carotenoid color variation in male contestcompetition. Using two experiments, we investigated the statussignaling function of the variable (ranging from yellow tored) carotenoid throat patch (collar) in the polygynous, sexually dimorphic red-collared widowbird (Euplectes ardens). First,we tested if the red collar functions as a dominance signalby painting spectrometrically controlled collar patches ontothe brown plumage of nonbreeding males and staging dyadic malecontests over food resources. Red-collared males dominatedorange males, which in turn dominated the control brown andnovel blue collars. Red dominance persisted when the collar manipulations were reversed within dyads and also when testedagainst testosterone implanted males. In the second experimentthe collar size and color of breeding males were manipulatedin the field before and after territories were established.All males with enlarged red and most with enlarged orange orreduced red collars obtained territories, whereas most maleswith reduced orange and all with blackened (removed) collarsfailed to establish or retain territories. In addition, amongthe territorial males, those with reduced signals defendedsmaller territories, received more intrusions, and spent moretime in aggressive interactions. Redness and, to a lesser extent,size of the carotenoid ornament both seem to independently indicate male dominance status or fighting ability in male contest competition.     

UNIDIRECTIONAL INTROGREESION OF A SEXUALLY SELECTED TRAIT ACROSS AN AVIAN HYBRID ZONE: A ROLE FOR FEMALE CHOICE?

Hybridization can be an evolutionary creative force by forming new polyploid species, creating novel genetic variation or acting as conduits of potentially advantageous traits between hybridizing forms. Evidence for the latter is often difficult to find because alleles under positive selection can spread rapidly across a hybrid zone and sweep to fixation. In Western Panama, an avian hybrid zone between two species of manakins in the genus Manacus exists where the unidirectional introgression of bright, yellow plumage into a white population provides evidence for the importance of hybrid zones as conduits of advantageous traits. Several lines of indirect evidence suggest that sexual selection favoring yellow plumage drives this asymmetrical spread, but more direct evidence is lacking. Along the edge of the hybrid zone, both yellow- and white-collared manakins are found in the same mating arenas or leks and compete for the same females ("mixed leks"), providing us with a unique opportunity to understand the dynamics of yellow plumage introgression. We studied these mixed leks to determine whether yellow males have a mating advantage over white males and, if so, whether the mating advantage is driven by male-male interactions, female choice, or both. We found that yellow males mated more than white males, suggesting that sexual selection favoring yellow males can, indeed, explain the spread of yellow plumage. However, we found that this advantage occurred only in mixed leks where the frequency of yellow males is greater than white males. This suggests that the advantage of yellow males may depend on the presence of other yellow males, which may slow the rate of introgression in leks where yellow frequency is low such as in areas where yellow males are beginning to colonize the white population. This, along with the geographic barrier posed by major rivers in the hybrid zone, may initially limit or slow the spread of yellow plumage. Finally, we found that yellow and white males were similar in aggression and body size, and held comparable positions within leks. Because these traits or factors are often important in or dictated by aggressive male-male interactions, these comparisons indicate that male-male interaction is not the primary mechanism for the spread of yellow plumage. However, white and yellow males received similar numbers of courtship visits from females but differed in the number of matings, suggesting that females actively rejected white in favor of yellow males. Our results indicate that sexual selection by female choice has driven the unidirectional introgression of yellow plumage into the white population, providing a mechanism for how hybrid zones act as conduits of novel and advantageous traits.     

APOA5 polymorphisms influence plasma triglycerides in young, healthy African Americans and whites of the CARDIA Study

Genetic variation in the apolipoprotein A-V gene (APOA5) has been associated with variation in plasma triglyceride (TG) levels in African American and white females and males older than 40 years and/or at increased risk of coronary artery disease. We have examined whether plasma TG levels are associated with 16 APOA5 polymorphisms in young (18-30 years) African American (1,075 females and 783 males) and white (1,041 females and 932 males) individuals of the Coronary Artery Risk Development in Young Adults (CARDIA) Study selected without regard to health. Plasma TG was significantly (P < 0.01) associated with markers 27376 and 28837 (-3A/G) in both white females and males, with 27709 (-1131T/C) and 29085 in white males, with 29009 (S19W) in African American females and white males, and with 30966 in African American females. No statistically significant associations were observed in African American males. These six single-nucleotide polymorphisms individually accounted for 0-0.78% of lnTG variation among white females, 0-2.46% among white males, and 0-0.69% among African American females. The results of our study suggest a small but replicable context-dependent influence of the APOA5 gene region on plasma TG levels in young, healthy individuals.     

Expression of cis-regulatory mutations of the white locus in metafemales of Drosophila melanogaster.

At the white eye colour locus, there are a number of alleles that have altered expression between males and females. To test these regulatory mutations of the white eye colour locus for their phenotypic expression in metafemales (3X; 2A) compared to diploid females and males, eleven alleles or transduced copies of white were analysed. Two alleles that exhibit dosage compensation between males and females (apricot, blood) also exhibit dosage compensation in metafemales. White-ivory and white-eosin, which fail to dosage compensate in males compared to females, but that are distinct physical lesions, also show a dosage effect in metafemales. Two alleles with greater expression in males than females (spotted, spotted-55) exhibit even lower expression in metafemales. Lastly, five transduced copies of white carrying three different lengths of the white promoter, but that all exhibit higher expression in males, show reduced expression in metafemales, exhibiting an inverse correlation between the level of expression and the dosage of the X chromosome. Because these alleles of white respond to dosage compensation in metafemales as a continuum of the male and female responses, it is concluded that the same basic mechanism of dosage compensation is involved and that the dosage of the X chromosome conditions the sexually dimorphic expression.     

Melanin-based coloration is a nondirectionally selected sex-specific signal of offspring development in the alpine swift

Two mutually exclusive hypotheses have been put forward to explain the evolution and adaptive function of melanin-based color traits. According to sexual selection theory melanism is a directionally selected signal of individual quality, whereas theory on the maintenance of genetic polymorphism proposes that alternative melanin-based variants achieve equal fitness. Alpine swift (Apus melba) males and females have a conspicuous patch of white feathers on the breast with their rachis varying continuously from white to black, and hence the breast varies from white to striated. If this trait is a sexually selected signal of quality, its expression should be condition dependent and the degree of melanism directionally selected. If variation in melanism is a polymorphism, its expression should be genetically determined and fitness of melanin-based variants equal. We experimentally tested these predictions by exchanging eggs or hatchlings between randomly chosen nests and by estimating survival and reproduction in relation to melanism. We found that breast melanism is heritable and that the environment and body condition do not significantly influence its expression. Between 5 and 50 days of age nestlings were heavier and their wings longer when breast feathers of their biological father were blacker, and they also fledged at a younger age. This shows that aspects of offspring quality covary positively with the degree of melanism. However, this did not result in directional selection because nestling survival and recruitment in the local breeding population were not associated with father breast melanism. Furthermore, adult survival, age at first reproduction and probability of skipping reproduction did not covary with the degree of melanism. Genetic variation in breast melanism is therefore maintained either because nonmelanic males achieve fitness similar to melanic males via a different route than producing fast-growing offspring, or because the advantage of producing fast-growing offspring is not sufficiently pronounced to result in directional selection.     

Timing of breeding and offspring number covary with plumage colour among Gyrfalcons Falco rusticolus

Plumage colour variation exists among Gyrfalcons throughout their Arctic and sub‐Arctic circumpolar distribution, ranging from white through silver and grey to almost black. Although different colour variants coexist within many populations, a few geographical regions, such as northern Greenland, possess a single variant, suggesting that local environments may influence plumage colour variation. In central‐west Greenland (66.5–67.5°N), where multiple colour variants exist, white male Gyrfalcons fathered significantly earlier clutches than grey males. No significant association was observed between female colour and lay date. However, significantly more offspring were produced by both male and female white Gyrfalcons than by grey variants when controlling for lay date, and silver Gyrfalcons produced an intermediate number of offspring for both sexes. This pattern was further supported by breeding plumage colour pairings. Grey females paired with grey males nested significantly later in the season and produced fewer offspring than those paired with white males, whereas no difference in lay date or offspring number was found between white males paired with white or with grey females. The difference in the number of offspring produced at each nest‐site was also inversely correlated with the distance to the nearest neighbouring nest, and grey males nested in closer proximity to other nests compared with white and silver colour variants. These results suggest that factors associated with territory occupancy and timing of breeding may regulate reproductive success differently between colour variants, with directional selection favouring light‐coloured Gyrfalcons and resulting in earlier lay date and a high frequency of white plumage colour variants in this population. Although gene flow exists between our study population and those further north (>75°N), white Gyrfalcons prevail where the breeding season duration is even shorter, suggesting that nesting chronology in combination with genetic drift may play an important role in influencing plumage colour polymorphism among Gyrfalcon populations.     

A label of health: a previous immune challenge is reflected in the expression of a female plumage trait

Studies of ornaments as indicators of individual quality have hitherto focused on males, while studies of female ornaments have been almost absent. However, females within the same species may show large variation in both outer appearance and individual quality. We experimentally examined the effect of an immune challenge on the size of white fringes in the wing feathers produced one to two months after the challenge in female common eiders (Somateria mollissima), a long-lived sea duck. Immune-challenged females produced feathers with smaller white fringes, showing that a previous immune challenge may affect the expression of a plumage trait. We also report an unexpected difference in the expression of this white plumage trait within the immune-challenged group, related to the specific immune response against one of the injected antigens.     

Red dominates black: agonistic signalling among head morphs in the colour polymorphic Gouldian finch

Recent sexual selection studies on the evolution of bird colouration have mainly focused on signals with a high level of condition-dependent variation, with much less attention given to colour traits whose expression is genetically controlled. Here, we experimentally tested the relative importance of a genetic colour polymorphism in determining male dominance in the Gouldian finch (Erythrura gouldiae), a species displaying three completely discrete but naturally co-occurring genetically inherited phenotypes; yellow-, red- (carotenoid) and black-headed (melanin) morphs. First, in staged dominance contests between unfamiliar birds of different head morphs, red-headed males dominated black-headed males, both of which dominated the yellow-headed birds. Second, within morphs, the intensity and size of the strongly ultraviolet-blue collar determined the outcome of these contests, and among the red-headed males, redder males dominated less chromatic birds. Lastly, when the dominance signal of red-headed birds was experimentally destabilized (i.e. blackened or reddened), naturally red-headed morphs continued to dominate both the black-and yellow-headed morphs. Together, these results suggest that intrinsic dominance-related behavioural differences between the three colour morphs, which are likely to influence the relative fitness of each morph, contribute to the complex selective patterns maintaining these three discrete phenotypes in relatively stable frequencies in wild populations.     

Sexual selection on plumage and behavior in an avian hybrid zone: experimental tests of male-male interactions

In western Panama, an unusual hybrid zone exists between white-collared manakins, Manacus candei, and golden-collared manakins, M. vitellinus. Unidirectional introgression of plumage traits from vitellinus into candei has created a region in which all definitively plumaged males have a collar that is lemon-colored. These males are nearly indistinguishable from white-collared candei genetically and morphometrically, but strongly resemble golden-collared vitellinus due to the introgression of secondary sexual plumage traits, particularly the lemon-colored collar. The introgression could be explained by sexual selection for golden-collared traits or by a series of mechanisms that do not invoke sexual selection (e.g., neutral diffusion, dominant allele). Sexual selection on male-male interactions implies behavioral differences among the plumage forms--specifically that golden- and lemon-collared males should be more aggressive than white-collared males. In contrast, the nonsexual hypotheses predict behavioral similarity between lemon- and white-collared males, based on their nearly identical genetics. We tested the sexual selection hypothesis experimentally, by presenting males with taxidermic mounts of the three forms. As response variables, we monitored vocalizations and attacks on the mounts by replicate subject males. Both golden-collared and lemon-collared males were more likely to attack than were white-collared males, as predicted under sexual selection but not by the nonsexual hypotheses. Lemon-collared males were more vocally reactive than either parental form, contrary to the prediction of the nonsexual hypotheses. Our study demonstrates that sexual selection on male-male interactions may play an important role in the dynamics of character evolution and hybrid zones.     

The adaptive significance of colour pattern polymorphism in the Australian scincid lizard Lampropholis delicata

Females of Lampropholis delicata are dimorphic for colour pattern, the difference between morphs being the presence or absence of a distinct white mid-lateral stripe. A less distinct striped morph occurs also in males. We evaluated alternative hypotheses for the maintenance of this polymorphism by examining temporal and spatial variation in morph frequency, testing for differential selection among morphs using data on body size and reproductive traits from preserved specimens, and experimentally manipulating colour pattern in free-ranging lizards of both sexes, to assess the influence of the lateral stripe on survival rates. We found that the relative frequency of striped individuals varied among populations and decreased from north to south in both sexes, coincident with an increasing incidence of regenerated tails. Morph frequencies did not change through time within a population. Striped gravid females appeared to survive better and produced larger clutches than did non-striped females. In our experimental study, the relationship between survival and colour morph differed between the two sexes; males painted with a white lateral stripe had lower survival than control (brown stripe) males, but survival did not differ between striped and control females. The different response in the two sexes may be due partly to differences in temperature and microhabitat selection. We propose that the white lateral stripe decreases susceptibility to predators in gravid females but increases risk of predation in males, especially in combination with low temperatures. The polymorphism might be maintained by: (1) opposing fitness consequences of the stripe in males and females; (2) sex-specific habitat selection; and (3) gene flow in combination with spatial variation in relative fitness of the two morphs.     

Is Midlife Occupational Physical Activity Related to Disability in Old Age? The SNAC-Kungsholmen Study

Objectives

Leisure-time physical activity (PA) has been established to be related to more years lived without disability. However, less is known about the relationship between occupational PA and disability in old age. The aim of the study was 1) to investigate whether midlife occupational PA is related to late-life disability, and 2) to test the hypothesis that the association differs according to the occupational categories of blue and white collar work.

Methods

The study population was derived from the Swedish National Study on Aging and Care, and consisted of a random sample of 1804 subjects aged 72 and above. The association of occupational PA during the longest held occupation with disability in old age was determined using logistic regression.

Results

There was no significant relationship between occupational PA and disability in personal or instrumental activities of daily living (ADL) after controlling for demographic and health-related factors. However, in stratified analyses moderate levels of occupational PA was associated with a lower odds ratio of dependency in personal ADL amongst white collar workers, compared to low level of occupational PA (OR = 0.34 95% C1 0.12–0.98).

Conclusions

Moderate levels of midlife occupational PA were associated with a decreased risk of personal ADL disability in old age among white collar workers, but not among blue collar workers. Our results highlight the importance of encouraging white collar workers to engage in physical activity during or outside work hours.     

The adaptive significance of electrophoretic mobility in phosphoglucose isomerase (PGI)

Sexual dimorphism in size and plumage was investigated in the lek-breeding Jackson's widowbird Euplectes jacksoni. Size dimorphism was evident from the sexual difference in tarsus length, reflecting skeletal body size. The long tail and the wide collar of breeding plumaged males were obvious secondary sex traits, but the wing was also longer than in subadults and might likewise be sexually selected (i.e. not merely a byproduct of body size). Possibly, the increased wingspan functions to save energy during the jump display. To reveal intrasexual selection, courtholders were compared with floaters (nuptial males captured off the leks), and were found to be larger in body size, wing length, collar and tail. The roles of inter-and intrasexual selection are discussed with reference to previously demonstrated female choice, and new results indicating competition for access to leks: quick reoccupation of deserted display courts, frequent observations of intruders and a finding that central males were more aggressive than peripherals. Observations imply that the collar is an agonistic signal that is neutral with respect to female choice. Possible contributions of natural selection to the sexual dimorphism are also considered.     

Genes regulated by mating, sperm, or seminal proteins in mated female Drosophila melanogaster

In Drosophila melanogaster, sperm and accessory gland proteins ("Acps," a major component of seminal fluid) transferred by males during mating trigger many physiological and behavioral changes in females (reviewed in ). Determining the genetic changes triggered in females by male-derived molecules and cells is a crucial first step in understanding female responses to mating and the female's role in postcopulatory processes such as sperm competition, cryptic female choice, and sexually antagonistic coevolution. We used oligonucleotide microarrays to compare gene expression in D. melanogaster females that were either virgin, mated to normal males, mated to males lacking sperm, or mated to males lacking both sperm and Acps. Expression of up to 1783 genes changed as a result of mating, most less than 2-fold. Of these, 549 genes were regulated by the receipt of sperm and 160 as a result of Acps that females received from their mates. The remaining genes whose expression levels changed were modulated by nonsperm/non-Acp aspects of mating. The mating-dependent genes that we have identified contribute to many biological processes including metabolism, immune defense, and protein modification.     

High -resolution scanning electron-microscopic studies on the three-dimensional structure of mitochondria and sarcoplasmic reticulum in the different twitch muscle fibers of the frog

Summary The three-dimensional structure of the mitochondria and sarcoplasmic reticulum (SR) in the three types of twitch fibers, i.e., the red, white and intermediate skeletal muscle fibers, of the vastus lateralis muscle of the Japanese meadow frog (Rana nigromaculata nigromaculata Hallowell) was examined by high resolution scanning electron microscopy, after removal of the cytoplasmic matrices.The small red fibers have numerous mitochondrial columns of large diameter, while the large white fibers have a small number of mitochondrial columns of small diameter. In the medium-size intermediate fibers, the number and diameter of the mitochondrial columns are intermediate between those of the red and white fibers.In all three types of fibers, the terminal cisternae and transverse tubules form triads at the level of each Z-line. The thick terminal cisternae continue into much thinner flat intermediate cisternae, through a transitional part where a row of tiny indentations can be observed. Numerous slender longitudinal tubules originating from the intermediate cisternae, extend longitudinally or obliquely and form elongated oval networks of various sizes in front of the A-band, then fuse to form the H-band collar (fenestrated collar) around the myofibrils. On the surface of the H-band collar, small fenestrations as well as tiny hollows are seen. The three-dimensional structure of SR is basically the same in all three muscle fiber-types. However, the SR is sparse on the surface of mitochondria, so the mitochondria-rich red fiber has a smaller total volume of SR than the mitochondria-poor white fiber. The volume of SR of the intermediate fiber is intermediate between other the two.     

Estimating the strength of sexual selection from Y-chromosome and mitochondrial DNA diversity

We show that a sex difference in the opportunity for selection results in sex differences in the strength of random genetic drift and thus creates different patterns of genetic diversity for maternally and paternally inherited haploid genes. We derive the effective population size Ne for a male-limited or female-limited haploid gene in terms of I, the "opportunity for selection" or the variance in relative fitness. Because the variance in relative fitness of males can be an order of magnitude larger than that of females, the Ne is much smaller for males than it is for females. We derive both nonequilibrium and equilibrium expressions for F(ST) in terms of I and show how the portion of I owing to sexual selection, Imates, that is, the variation among males in mate numbers, is a simple function of the F's for cytoplasmic (female inherited) and Y-linked (male inherited) genes. Because multiple, transgenerational data are lacking to apply the nonequilibrium expression, we apply only the equilibrium model to published data on Y chromosome and mitochondrial sequence divergence in Homo sapiens to quantify the opportunity for sexual selection. The estimate suggests that sexual selection in humans represents a minimum of 54.8% of total selection, supporting Darwin's proposal that sexual selection has played a significant role in human evolution and the recent proposal regarding a shift from polygamy to monogamy in humans.     

Genome-wide germline-enriched and sex-biased expression profiles in Caenorhabditis elegans

We performed a genome-wide analysis of gene expression in C. elegans to identify germline- and sex-regulated genes. Using mutants that cause defects in germ cell proliferation or gametogenesis, we identified sets of genes with germline-enriched expression in either hermaphrodites or males, or in both sexes. Additionally, we compared gene expression profiles between males and hermaphrodites lacking germline tissue to define genes with sex-biased expression in terminally differentiated somatic tissues. Cross-referencing hermaphrodite germline and somatic gene sets with in situ hybridization data demonstrates that the vast majority of these genes have appropriate spatial expression patterns. Additionally, we examined gene expression at multiple times during wild-type germline development to define temporal expression profiles for these genes. Sex- and germline-regulated genes have a non-random distribution in the genome, with especially strong biases for and against the X chromosome. Comparison with data from large-scale RNAi screens demonstrates that genes expressed in the oogenic germline display visible phenotypes more frequently than expected.     

Determinants of age-dependent change in a secondary sexual character

Many secondary sexual characters vary in a systematic way with the age of individuals, with young and old individuals displaying at lower levels than individuals of intermediate age. Analyses quantifying the within-individual and among-individual components of phenotypic variation can help partition effects of phenotypic plasticity and selective mortality. We analysed phenotypic variation in the expression of a secondary sexual character, tail length, in male and female barn swallows Hirundo rustica from four European populations studied during 11-26 years, using linear mixed effect models to describe age-related expression. Tail length increased from yearlings to intermediate aged birds with a subsequent decrease at old age. In males, this age-related pattern was because of both within-subject and between-subject effects, with no difference among populations. Males having longer lifespan had shorter tails when young than those having shorter lifespan. Females showed similar patterns of age-related variation as males, with no difference among populations. The major difference between sexes was that the between-subject effects (i.e. disappearance effects or selection) were much more important for males compared to females for which lifetime variation in tail length was mainly because of a within-subject effect (i.e., a plastic response). These findings suggest that whereas males trade greater expression of the secondary sexual character at young age against longevity, that was not the case for females. This is consistent with tail length being more costly in males than in females, with the cost of long tails potentially being offset by elevated mating success, whereas that is not the case in females.     

Sexual selection of multiple handicaps in the red-collared widowbird: female choice of tail length but not carotenoid display

Although sexual selection through female choice explains exaggerated male ornaments in many species, the evolution of the multicomponent nature of most sexual displays remains poorly understood. Theoretical models suggest that handicap signaling should converge on a single most informative quality indicator, whereas additional signals are more likely to be arbitrary Fisherian traits, amplifiers, or exploitations of receiver psychology. Male nuptial plumage in the highly polygynous red-collared widowbird (Euplectes ardens) comprises two of the commonly advocated quality advertisements (handicaps) in birds: a long graduated tail and red carotenoid coloration. Here we use multivariate selection analysis to investigate female choice in relation to male tail length, color (reflectance) of the collar, other aspects of morphology, ectoparasite load, display rate, and territory quality. The order and total number of active nests obtained are used as measures of male reproductive success. We demonstrate a strong female preference and net sexual selection for long tails, but marginal or no effects of color, morphology, or territory quality. Tail length explained 47% of male reproductive success, an unusually strong fitness effect of natural ornament variation. Fluctuating tail asymmetry was unrelated to tail length, and had no impact on mating success. For the red collar, there was negative net selection on collar area, presumably via its negative relationship with tail length. None of the color variables (hue, chroma, and brightness) had significant selection differentials, but a partial effect (selection gradient) of chroma might represent a color preference when tail length is controlled for. We suggest that the red collar functions in male agonistic interactions, which has been strongly supported by subsequent work. Thus, female choice targets only one handicap, extreme tail elongation, disregarding or even selecting against the carotenoid display. We discuss whether long tails might be better indicators of genetic quality than carotenoid pigmentation. As regards the evolution of multiple ornaments, we propose that multiple handicap signaling is stable not because of multiple messages but because of multiple receivers, in this case females and males.     

The interplay of light and the circadian clock. Independent dual regulation of clock-controlled gene ccg-2(eas).

Ambient light is the major agent mediating entrainment of circadian rhythms and is also a major factor influencing development and morphogenesis. We show that in Neurospora crassa the expression of clock-controlled gene 2 (ccg-2), a gene under the control of the circadian clock and allelic to the developmental gene easy wettable (eas), is regulated by light in wild-type strains. Light elicits a direct and important physiological effect on ccg-2(eas) expression as demonstrated using several mutant Neurospora strains. In white collar mutants (wc-1 and wc-2) that are "blind" to blue light, ccg-2(eas) mRNA shows no variation following illumination with saturating light. By contrast, ccg-2(eas) mRNA is photoinduced in clock-null strains such as frequency (bd;frq). The results in the clock mutants show that an intact circadian oscillator is not required for light induction of ccg-2(eas). Thus, ccg-2(eas) is subject to a dual regulation that involves separable regulation by light and circadian rhythm.