Social preferences of domestic hens for domestic vs. red junglefowl males and females

The striking dimorphism of male and female junglefowl may have evolved in large part due to female choice of mates. However, domestic fowl hens consistently chose domestic, not junglefowl, males as well as females in preference tests. Either genetic predisposition to favor junglefowl morphological features over those of the domestic types utilized in these tests was overridden by experience or such genetic predisposition does not exist in the breed we tested.     

The effects of age and previous experience on social rank in female red junglefowl, Gallus gallus spadiceus

Social rank can influence lifetime reproductive success and therefore fitness. We examined the effects of morphology, age, previous social experience and aggressiveness on social rank in all-female flocks of red junglefowl. None of the morphological characters measured (mass, tarsus length, comb height or comb length) appeared to play a role in determining rank. Older females were not more likely to be dominant, while previous social experience and aggression levels were both important in dominance determination. Flock-experienced hens were more likely to be dominant as were more aggressive individuals. Red junglefowl females most likely use a combination of characters to establish social order in a newly formed flock. Copyright 2000 The Association for the Study of Animal Behaviour.     

中国家鸡的起源探讨

测定了6个家鸡品种30个个体的线粒体D0环区539bp的碱基序列,并与GenBank中的红原鸡、灰原鸡、绿原鸡、黑尾原鸡及鹌鹑的相应序列作比较分析,构建了分析系统树。结果表明,原鸡属4个种间差异较大,其中家鸡与泰国及邻近地区红原鸡关系最近,与印尼红原鸡、黑尾原鸡、灰原鸡、绿原鸡及鹌鹑的关系依次变远。提示中国家鸡可能起源于泰国及邻近地区的红原鸡。结果还显示,该原鸡的两个亚种（G.g.gallus和G.g.spadiceus)间的遗传分化程度显著低于家鸡亚种内的分化程度,这两个亚种似乎可以归为同一亚种。     

Sex-specific, counteracting responses to inbreeding in a bird

Inbreeding often depresses offspring fitness. Because females invest more than males in a reproductive event, inbreeding is expected to be more costly to mothers than fathers, creating a divergence between the reproductive interests of each sex and promoting sex-specific inbreeding strategies. Males and females may bias the probability of inbreeding by selecting copulation partners, and, in sexually promiscuous species, through male strategic sperm investment in different females and female selection of the sperm of different males. However, these processes are often difficult to study, and the way that different male and female strategies interact to determine inbreeding remains poorly understood. Here we demonstrate sex-specific, counteracting responses to inbreeding in the promiscuous red junglefowl, Gallus gallus. First, a male was just as likely to copulate with his full-sib sister as with an unrelated female. In addition, males displayed a tendency to: (i) initiate copulation faster when exposed to an unrelated female than when exposed to a sister, and (ii) inseminate more sperm into sisters than into unrelated females. Second, females retained fewer sperm following insemination by brothers, thus reducing the risk of inbreeding and counteracting male inbreeding strategies.     

The origin and genetic diversity of Chinese native chicken breeds

The first 539 bases of mitochondrial DNA D-loop region of six Chinese native chicken breeds (Gallus gallus domesticus) were sequenced and compared to those of the red junglefowl (Gallus gallus), the gray junglefowl (Gallus sonneratii), the green junglefowl (Gallus varius) and Lafayette's junglefowl (Gallus lafayettei) reported in GenBank, and the phylogenetic trees for the chickens were constructed based on the D-loop sequences. The results showed that the four species of the genus Gallus had great differences among each other, the G. g. domesticus was closest to the red junglefowl in Thailand and its adjacent regions, suggesting the Chinese domestic fowl probably originated from the red junglefowl in these regions. The two subs pecies of Thailand, G. g. gallus and G. g. spadiceus, should belong to one subspecies because of their resemblance. In the case of native breeds, there existed a great difference between the egg breeds and general purpose breeds, which suggested different maternal origins of the two types.     

Fertility Advertisement in Birds: a Means of Inciting Male-male Competition?

Certain loud calls made by female red junglefowl and Lapland longspurs are given most frequently immediately after egg laying, when a copulation should have the highest probability of fertilizing the next egg to be laid. In these species there is considerable male-male interaction for access to fertilizable females, and males are attracted to or follow females making these calls. Based on our interpretation of these calls, we develop a general hypothesis to predict the pattern of occurrence of fertility advertisement both within and among bird species. We hypothesize that certain loud calls given by female birds before and during the egg-laying period are designed to advertise fertility and thereby incite male-male competition. This hypothesis predicts that calls advertising female fertility should most often occur soon after an egg is laid (i.e. during the period of highest fertility) but may also occur at any time during the female's fertilizable period. Such calls are unlikely to be given by females in strictly monogamous species (especially those with long-term pair bonds) because in these species each female usually mates with only a single male and extra-pair copulations are avoided. Although reports of loud female calls associated with copulation are rare in the literature, the 18 examples we found (including junglefowl and longspurs) were predominantly (15/18) in species adopting mating systems other than strict monogamy, and these calls were most commonly and disproportionately reported in multi-male mating systems. This form of “estrus” in birds may be widespread because few species appear to be strictly monogamous, but it will be difficult to study because the period of high fertility for female birds is so short.     

Cryptic preference for MHC-dissimilar females in male red junglefowl,Gallus gallus

An increasing number of studies test the idea that females increase offspring fitness by biasing fertilization in favour of genetically compatible partners; however, few have investigated or controlled for corresponding preferences in males. Here, we experimentally test whether male red junglefowl, Gallus gallus, prefer genetically compatible females, measured by similarity at the major histocompatibility complex (MHC), a key gene complex in vertebrate immune function. Theory predicts that because some degree of MHC heterozygosity favours viability, individuals should prefer partners that carry MHC alleles different from their own. While male fowl showed no preference when simultaneously presented with an MHC-similar and an MHC-dissimilar female, they showed a ‘cryptic’ preference, by allocating more sperm to the most MHC-dissimilar of two sequentially presented females. These results provide the first experimental evidence that males might respond to the MHC similarity of a female through differential ejaculate expenditure. By revealing that cryptic male behaviours may bias fertilization success in favour of genetically compatible partners, this study demonstrates the need to experimentally disentangle male and female effects when studying preferences for genetically compatible partners.     

Quantitative genetics of sexually dimorphic traits and capture of genetic variance by a sexually-selected condition-dependent ornament in red junglefowl (Gallus gallus)

We studied the quantitative genetics of sexually selected traits in a captive population of red junglefowl (Gallus gallus L.) using a multi-generational 'animal model' approach. We found significant heritability of mass, tarsus length (both strongly sexually dimorphic), residual mass, and male comb (a fleshy head ornament) length. Residual mass has a genetic correlation between the sexes smaller than unity and so could show partially independent responses to selection in the two sexes. In males, tarsus length and mass were not genetically correlated, and this produced a negative genetic correlation between tarsus length and residual mass. The male red junglefowl's comb, an ornament influencing female choice, is highly condition dependent. We show that expression of this ornament is heritable, however, and shows strong genetic correlation with a condition index, residual mass. Because residual mass is partly influenced by various aspects of condition, it appears that comb size has 'captured' genetic variability in condition.     

Multiple aspects of condition influence a heritable sexual trait: a synthesis of the evidence for capture of genetic variance in red junglefowl

A long‐standing question in the study of sexual selection is: if an attractive male trait signals heritable aspects of quality, how does genetic variation associated with that trait persist in the face of directional selection? It has been proposed that sensitivity to condition in general could allow a sexual signal to show heritable variation even under persistent directional selection. The multitude of genes underlying components of condition present a large target for mutation, and so genetic variance in condition may not be readily exhausted by selection. Insights gained in studies of the red junglefowl are relevant to this hypothesis, and provide a model for research in other systems. The fleshy comb of male red junglefowl is among the best‐studied sexual signals. Multiple components of condition, such as health, immune function, testosterone, and social status, influence comb growth, and, in the present study, we provide the first comprehensive integration of this large body of evidence to lay out the compelling case for condition‐dependence of the junglefowl comb. Variation in comb size is heritable and, most important to this issue, is genetically correlated with heritable variation in a body condition index. Although understanding of the red junglefowl's comb is far from complete, it exceeds our understanding of many other sexual signals and provides an empirical model for the study of condition‐dependent signals in general. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 92 , 651–660.     

Social mediation of sexually selected ornamentation and steroid hormone levels in male junglefowl

Honest signals in sexual selection may be maintained by a variety of mechanisms. Comb size in male red junglefowl, Gallus gallus, a well-known predictor of female mate choice, is mediated by health and condition. Social status has also been shown to mediate comb size. To determine whether hormones related to male dominance behaviour might be related to social status and comb size, we monitored changes in plasma levels of testosterone and corticosterone following manipulation of social status. We removed young adult male junglefowl from their all-male flocks and placed them either in individual cages or in smaller flocks. We measured comb size and other morphological variables, as well as testosterone and corticosterone levels, before and after the manipulation of social status. Males in small flocks showed reduced comb growth, increased corticosterone and decreased testosterone levels compared with individually housed males. Within flocks, comb size was positively related to dominance rank and to testosterone level, although not always significantly. However, dominance rank was unrelated to either corticosterone or testosterone levels. In isolated birds, testosterone level was not related to comb size. Corticosterone level was not related to comb size in either treatment group. Our results are consistent with social mediation of comb size and testosterone and corticosterone levels, but it is unclear whether these hormones contribute to the observed social limitation of comb length.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved .     

Chicken domestication: an updated perspective based on mitochondrial genomes

Domestic chickens (Gallus gallus domesticus) fulfill various roles ranging from food and entertainment to religion and ornamentation. To survey its genetic diversity and trace the history of domestication, we investigated a total of 4938 mitochondrial DNA (mtDNA) fragments including 2843 previously published and 2095 de novo units from 2044 domestic chickens and 51 red junglefowl (Gallus gallus). To obtain the highest possible level of molecular resolution, 50 representative samples were further selected for total mtDNA genome sequencing. A fine-gained mtDNA phylogeny was investigated by defining haplogroups A–I and W–Z. Common haplogroups A–G were shared by domestic chickens and red junglefowl. Rare haplogroups H–I and W–Z were specific to domestic chickens and red junglefowl, respectively. We re-evaluated the global mtDNA profiles of chickens. The geographic distribution for each of major haplogroups was examined. Our results revealed new complexities of history in chicken domestication because in the phylogeny lineages from the red junglefowl were mingled with those of the domestic chickens. Several local domestication events in South Asia, Southwest China and Southeast Asia were identified. The assessment of chicken mtDNA data also facilitated our understanding about the Austronesian settlement in the Pacific.     

The measure and significance of Bateman's principles

Bateman''s principles explain sex roles and sexual dimorphism through sex-specific variance in mating success, reproductive success and their relationships within sexes (Bateman gradients). Empirical tests of these principles, however, have come under intense scrutiny. Here, we experimentally show that in replicate groups of red junglefowl, Gallus gallus, mating and reproductive successes were more variable in males than in females, resulting in a steeper male Bateman gradient, consistent with Bateman''s principles. However, we use novel quantitative techniques to reveal that current methods typically overestimate Bateman''s principles because they (i) infer mating success indirectly from offspring parentage, and thus miss matings that fail to result in fertilization, and (ii) measure Bateman gradients through the univariate regression of reproductive over mating success, without considering the substantial influence of other components of male reproductive success, namely female fecundity and paternity share. We also find a significant female Bateman gradient but show that this likely emerges as spurious consequences of male preference for fecund females, emphasizing the need for experimental approaches to establish the causal relationship between reproductive and mating success. While providing qualitative support for Bateman''s principles, our study demonstrates how current approaches can generate a misleading view of sex differences and roles.     

Effects of the domestic thyroid stimulating hormone receptor (TSHR) variant on the hypothalamic-pituitary-thyroid axis and behavior in chicken

Domestic chickens are less fearful, have a faster sexual development, grow bigger, and lay more eggs than their primary ancestor, the red junglefowl. Several candidate genetic variants selected during domestication have been identified, but only a few studies have directly linked them with distinct phenotypic traits. Notably, a variant of the thyroid stimulating hormone receptor (TSHR) gene has been under strong positive selection over the past millennium, but it’s function and mechanisms of action are still largely unresolved. We therefore assessed the abundance of the domestic TSHR variant and possible genomic selection signatures in an extensive data set comprising multiple commercial and village chicken populations as well as wild-living extant members of the genus Gallus. Furthermore, by mean of extensive backcrossing we introgressed the wild-type TSHR variant from red junglefowl into domestic White Leghorn chickens and investigated gene expression, hormone levels, cold adaptation, and behavior in chickens possessing either the wild-type or domestic TSHR variant. While the domestic TSHR was the most common variant in all studied domestic populations and in one of two red junglefowl population, it was not detected in the other Gallus species. Functionally, the individuals with the domestic TSHR variant had a lower expression of the TSHR in the hypothalamus and marginally higher in the thyroid gland than wild-type TSHR individuals. Expression of TSHB and DIO2, two regulators of sexual maturity and reproduction in birds, was higher in the pituitary gland of the domestic-variant chickens. Furthermore, the domestic variant was associated with higher activity in the open field test. Our findings confirm that the spread of the domestic TSHR variant is limited to domesticated chickens, and to a lesser extent, their wild counterpart, the red junglefowl. Furthermore, we showed that effects of genetic variability in TSHR mirror key differences in gene expression and behavior previously described between the red junglefowl and domestic chicken.     

Domestication effects on behavioural synchronization and individual distances in chickens (Gallus gallus)

Behavioural synchrony (allelomimetic behaviour), and inter-individual distances are aspects of social and anti-predator strategies which may have been affected by domestication. Chickens are known to adjust synchronization and inter-individual distances depending on behaviour. We hypothesized that White Leghorn (WL) chickens would show less synchronized behaviour than the ancestor, the red jungle fowl (RJF). Sixty birds, 15 female and 15 male WL and the same number of RJF (28 weeks old) were studied in groups of three in furnished pens (1 m × 2 m) for 24 consecutive hours per group, following 24 h of habituation. Video tapes covering 4 h per group (dawn, 9-10 am, 1-2 pm and dusk) were analysed. Red junglefowl perched significantly more, but there were no breed effects on the frequency or daily rhythm of any other activities, or on average inter-individual distances. Red junglefowl were more synchronized during perching and a tendency for the same was found for social behaviour. After performance of the two most synchronized behaviours, perching and comfort behaviour, individual distance increased more for RJF than WL. According to this study domestication of chickens appears not to have significantly altered the relative frequencies of different activities or average inter-individual distances, but have caused some changes in behavioural synchronization and maintenance of activity-specific inter-individual distances in chickens. The changes may indicate an adaptive response to captivity and domestication.     

Asymmetries,heterosis, and phenotypic profiles of red junglefowl,White Plymouth Rocks,and F<Subscript>1</Subscript> and F<Subscript>2</Subscript> reciprocal crosses

During the domestication of farm animals, humans have manipulated genetic variation for growth and reproduction through artificial selection. Here, data are presented for growth, reproductive, and behavior traits for the red junglefowl, a line of White Plymouth Rock chickens, and their F₁ and F₂ reciprocal crosses. Intra- and intergenerational comparisons for growth related traits reflected considerable additive genetic variation. In contrast, those traits associated with reproduction exhibited heterosis. The role of sexual selection was seen in the evolution of prominent secondary sexual ornaments that lend to female choice and male-male competition. The large differences between parental lines in fearfulness to humans were only mitigated slightly in the intercross generations. Whereas, overall F₁ generation heterosis was not transferred to the F₂, there was developmental stability in the F₂, as measured by relative asymmetry of bilateral traits. Through multigenerational analyses between the red junglefowl and the domestic White Plymouth Rocks, we observed plasticity and considerable residual genetic variation. These factors likely facilitated the adaptability of the chicken to a broad range of husbandry practices throughout the world.     

The contrasting role of male relatedness in different mechanisms of sexual selection in red junglefowl

In structured populations, competition for reproductive opportunities should be relaxed among related males. The few tests of this prediction often neglect the fact that sexual selection acts through multiple mechanisms, both before and after mating. We performed experiments to study the role of within‐group male relatedness across pre‐ and postcopulatory mechanisms of sexual selection in social groups of red junglefowl, Gallus gallus, in which two related males and one unrelated male competed over females unrelated to all the males. We confirm theoretical expectations that, after controlling for male social status, competition over mating was reduced among related males. However, this effect was contrasted by other sexual selection mechanisms. First, females biased male mating in favor of the unrelated male, and might also favor his inseminations after mating. Second, males invested more—rather than fewer—sperm in postcopulatory competition with relatives. A number of factors may contribute to explain this counterintuitive pattern of sperm allocation, including trade‐offs between male investment in pre‐ versus postcopulatory competition, differences in the relative relatedness of pre‐ versus postcopulatory competitors, and female bias in sperm utilization in response to male relatedness. Collectively, these results reveal that within‐group male relatedness may have contrasting effects in different mechanisms of sexual selection.     

Cryptic female choice favours sperm from major histocompatibility complex-dissimilar males

Cryptic female choice may enable polyandrous females to avoid inbreeding or bias offspring variability at key loci after mating. However, the role of these genetic benefits in cryptic female choice remains poorly understood. Female red junglefowl, Gallus gallus, bias sperm use in favour of unrelated males. Here, we experimentally investigate whether this bias is driven by relatedness per se, or by similarity at the major histocompatibility complex (MHC), genes central to vertebrate acquired immunity, where polymorphism is critical to an individual''s ability to combat pathogens. Through experimentally controlled natural matings, we confirm that selection against related males'' sperm occurs within the female reproductive tract but demonstrate that this is more accurately predicted by MHC similarity: controlling for relatedness per se, more sperm reached the eggs when partners were MHC-dissimilar. Importantly, this effect appeared largely owing to similarity at a single MHC locus (class I minor). Further, the effect of MHC similarity was lost following artificial insemination, suggesting that male phenotypic cues might be required for females to select sperm differentially. These results indicate that postmating mechanisms that reduce inbreeding may do so as a consequence of more specific strategies of cryptic female choice promoting MHC diversity in offspring.     

Antlers honestly advertise sperm production and quality

Evolutionary theory proposes that exaggerated male traits have evolved via sexual selection, either through female mate choice or male-male competition. While female preferences for ornamented males have been amply demonstrated in other taxa, among mammals sexual characters are commonly regarded as weapons whose main function is to enhance male competitiveness in agonistic encounters. One particularly controversial hypothesis to explain the function of male sexual characters proposes that they advertise male fertility. We test this hypothesis in red deer (Cervus elaphus), a species where sexual characters (antlers) reach an extreme degree of elaboration. We find that a global measure of relative antler size and complexity is associated with relative testes size and sperm velocity. Our results exclude the possibility that condition dependence, age or time of culling, drive these associations. Red deer antlers could signal male fertility to females, the ability to avoid sperm depletion throughout the reproductive season and/or the competitive ability of ejaculates. By contrast, male antlers could also signal to other males not only their competitive ability at the behavioural level (fighting ability) but also at the physiological level (sperm competition).     

Quantitative genetics of ontogeny of sexual dimorphism in red junglefowl (Gallus gallus)

We studied phenotypic patterns and underlying quantitative genetics of development of sexual size dimorphism in red junglefowl (Gallus gallus). Using a multigenerational pedigree and the 'animal model' technique, we found significant heritability for many of the size and growth-related traits we examined, as well as significant genetic correlations among them. Despite sexual size dimorphism throughout posthatching ontogeny, the genetic correlation between males and females for all size measurements and growth parameters remained high. Significant positive phenotypic and genetic correlations between the fastest rate of growth and mass at week 26 (near asymptote) indicate that faster growth when young promotes larger adult size. However, age at which peak growth is reached does not appear to be phenotypically or genetically correlated with adult size. Positive genetic correlations within traits among ages were common, demonstrating that the genetic variance important to growth is relatively consistent among ages. However, male mass and tarsus length showed no genetic correlation between week 0 values and those from later ages. The body size traits of mass and tarsus length were genetically correlated with each other in females, but this pattern was not significant in males. Thus, despite striking sexual dimorphism in size and growth trajectories, size dimorphic traits in junglefowl show, with some exceptions, genetic integration between the sexes, among ages, and between traits.     

Pre‐ and postcopulatory sexual selection favor aggressive,young males in polyandrous groups of red junglefowl

A challenge in evolutionary biology is to understand the operation of sexual selection on males in polyandrous groups, where sexual selection occurs before and after mating. Here, we combine fine‐grained behavioral information (>41,000 interactions) with molecular parentage data to study sexual selection in replicated, age‐structured groups of polyandrous red junglefowl, Gallus gallus. Male reproductive success was determined by the number of females mated (precopulatory sexual selection) and his paternity share, which was driven by the polyandry of his female partners (postcopulatory sexual selection). Pre‐ and postcopulatory components of male reproductive success covaried positively; males with high mating success also had high paternity share. Two male phenotypes affected male pre‐ and postcopulatory performance: average aggressiveness toward rival males and age. Aggressive males mated with more females and more often with individual females, resulting in higher sexual exclusivity. Similarly, younger males mated with more females and more often with individual females, suffering less intense sperm competition than older males. Older males had a lower paternity share even allowing for their limited sexual exclusivity, indicating they may produce less competitive ejaculates. These results show that—in these populations—postcopulatory sexual selection reinforces precopulatory sexual selection, consistently promoting younger and more aggressive males.