Phenotypic variation and fluctuating asymmetry in sexually dimorphic feather ornaments in relation to sex and mating system

Secondary sexual characters have been hypothesized to demonstrate increased phenotypic variation between and within individuals as compared to ordinary morphological traits. We tested whether this was the case by studying phenotypic variation, expressed as the coefficient of variation (CV), and developmental instability, measured as fluctuating asymmetry (FA), in ornamental and non-ornamental traits of 70 bird species with feather ornamentation while controlling for similarity among species due to common descent. Secondary sexual characters differed from ordinary morphological traits by showing large phenotypic CV and FA. This difference can be explained by the different mode of selection operating on each kind of trait: a history of intense directional (ornaments) and stabilizing selection (non-ornaments). Phenotypic variation is reduced in the sex with more intense sexual selection (males), but does not differ among species with different mating systems. The strength of stabilizing selection arising from natural selection is associated with decreased CV (wing CV is smaller than tarsus or tail CVs). We found evidence of FA being reduced in ornamental feathers strongly affected by aerodynamics (tail feathers) compared to other ornaments, but only in females. In conclusion, CV and FA were not related, suggesting mat phenotypic plasticity and developmental instability are independent components of phenotypic variation.     

Genetics of body shape and armour variation in threespine sticklebacks

Patterns of genetic variation and covariation can influence the rate and direction of phenotypic evolution. We explored the possibility that the parallel morphological evolution seen in threespine stickleback (Gasterosteus aculeatus) populations colonizing freshwater environments is facilitated by patterns of genetic variation and covariation in the ancestral (marine) population. We estimated the genetic (G) and phenotypic (P) covariance matrices and directions of maximum additive genetic (g(max) ) and phenotypic (p(max) ) covariances of body shape and armour traits. Our results suggest a role for the ancestral G in explaining parallel morphological evolution in freshwater populations. We also found evidence of genetic constraints owing to the lack of variance in the ancestral G. Furthermore, strong genetic covariances and correlations among traits revealed that selective factors responsible for threespine stickleback body shape and armour divergence may be difficult to disentangle. The directions of g(max) and p(max) were correlated, but the correlations were not high enough to imply that phenotypic patterns of trait variation and covariation within populations are very informative of underlying genetic patterns.     

浙江楠种群表型变异

以探讨浙江楠(Phoebe chekiangensis)种群间和种群内的表型变异程度和变异规律为目的, 对其分布区9个种群的10个表型性状进行了研究。采用巢式方差分析、变异系数、多重比较和相关分析等多种分析方法, 对种群间和种群内的表型多样性及其与地理生态因子的相关性进行了讨论。研究结果表明: 浙江楠叶片、种子等表型性状在种群间和种群内均存在着丰富的变异, 10个性状在种群间、种群内的差异均达极显著水平; 种群内的变异(21.74%)大于种群间的变异(18.45%), 表型性状的平均分化系数为41.43%, 种群内变异是浙江楠表型变异的主要变异来源。各性状平均变异系数为12.78%, 变化幅度为6.50%-19.38%。种群间叶片的平均变异系数(16.99%)高于种群间种子的平均变异系数(8.58%), 表明种子的稳定性高于叶片。叶宽、叶面积与叶片其他性状间多呈显著或极显著相关, 种子千粒重与种子宽、种子体积显著正相关; 而种子和叶片的各性状间多无显著相关性。种子宽、种子体积和种子千粒重随着海拔的增加出现减小的趋势, 种子体积受纬度控制, 随着纬度的增大表现出增大的趋势。     

The evolution of male genitalia: patterns of genetic variation and covariation in the genital sclerites of the dung beetle Onthophagus taurus

Three main hypotheses, have been invoked to explain divergent genital evolution, the lock and key, pleiotropy, and sexual selection hypotheses, each of which make different predictions about how genital traits are inherited. Here we used a half-sib breeding design to examine the patterns of genetic variation and covariation between male genital sclerites, and their covariance with general body morphology in the dung beetle Onthophagus taurus. We found CV(A)'s and CV(P)'s were similar for both genital and general morphological traits and that CV(R)'s were large for both trait types. We found that male genital sclerites were negatively genetically correlated with general morphological traits. Variation in male genital morphology has direct implications for a male's fertilization success and the resulting sexual selection acting on male genitalia is predicted to maintain high levels of additive genetic variance. Contrary to this prediction, we found that individual genital sclerites all had low levels of additive genetic variance and large maternal and environmental sources of variation. Our data suggest that the genital sclerites in O. taurus are not inherited independently but as a genetically integrated unit. More importantly, the way the different sclerites function to influence male fertilization success reflects this genetic integration. Even though levels of V(A) in individual genital sclerites may be low, there may still be sufficient V(A) in multivariate trait space for selection to generate evolutionary change in the overall morphology of male genitalia.     

Comparing Mutational Variabilities

We have reviewed the available data on V(M), the amount of genetic variation in phenotypic traits produced each generation by mutation. We use these data to make several qualitative tests of the mutation-selection balance hypothesis for the maintenance of genetic variance (MSB). To compare V(M) values, we use three dimensionless quantities: mutational heritability, V(M)/V(E); the mutational coefficient of variation, CV(M); and the ratio of the standing genetic variance to V(M), V(G)/V(M). Since genetic coefficients of variation for life history traits are larger than those for morphological traits, we predict that under MSB, life history traits should also have larger CV(M). This is confirmed; life history traits have a median CV(M) value more than six times higher than that for morphological traits. V(G)/V(M) approximates the persistence time of mutations under MSB in an infinite population. In order for MSB to hold, V(G)/V(M) must be small, substantially less than 1000, and life history traits should have smaller values than morphological traits. V(G)/V(M) averages about 50 generations for life history traits and 100 generations for morphological traits. These observations are all consistent with the predictions of a mutation-selection balance model.     

The genetic basis of early‐life morphological traits and their relation to alternative male reproductive tactics in Atlantic salmon

Although heritability estimates for traits potentially under natural selection are increasingly being reported, their estimation remains a challenge if we are to understand the patterns of adaptive phenotypic change in nature. Given the potentially important role of selection on the early life phenotype, and thereby on future life history events in many fish species, we conducted a common garden experiment, using the Atlantic salmon (Salmo salar L.), with two major aims. The first objective is to determine how the site of origin, the paternal sexual tactic and additive genetic effects influence phenotypic variation of several morphological traits at hatching and emergence. The second aim is to test whether a link exists between phenotypic characteristics early in life and the incidence of male alternative tactics later in life. We found no evidence of a site or paternal effect on any morphological trait at hatching or emergence, suggesting that the spatial phenotypic differences observed in the natural river system from which these fish originated are mainly environmentally driven. However, we do find significant heritabilities and maternal effects for several traits, including body size. No direct evidence was found correlating the incidence of precocious maturation with early life characteristics. We suggest that under good growing conditions, body size and other traits at early developmental stages are not reliable cues for the surpassing of the threshold values associated with male sexual development.     

Patterns of genetic variation and covariation in ejaculate traits reveal potential evolutionary constraints in guppies

Ejaculates comprise multiple and potentially interacting traits that determine male fertility and sperm competitiveness. Consequently, selection on these traits is likely to be intense, but the efficacy of selection will depend critically on patterns of genetic variation and covariation underlying their expression. In this study, I provide a prospective quantitative genetic analysis of ejaculate traits in the guppy Poecilia reticulata, a highly promiscuous live-bearing fish. I used a standard paternal half-sibling breeding design to characterize patterns of genetic (co)variation in components of sperm length and in vitro sperm performance. All traits exhibited high levels of phenotypic and additive genetic variation, and in several cases, patterns of genetic variation was consistent with Y-linkage. There were also highly significant negative genetic correlations between the various measures of sperm length and sperm performance. In particular, the length of the sperm's midpiece was strongly, negatively and genetically correlated with sperm's swimming velocity-an important determinant of sperm competitiveness in this and other species. Other components of sperm length, including the flagellum and head, were independently and negatively genetically correlated with the proportion of live sperm in the ejaculate (sperm viability). Whether these relationships represent evolutionary trade-offs depends on the precise relationships between these traits and competitive fertilization rates, which have yet to be fully resolved in this (and indeed most) species. Nevertheless, these prospective analyses point to potential constraints on ejaculate evolution and may explain the high level of phenotypic variability in ejaculate traits in this species.     

Patterns of morphological and genetic variability in UK populations of the shore crab, Carcinus maenas Linnaeus, 1758 (Crustacea: Decapoda: Brachyura)

Previous research has identified extensive inter-population variability in the morphology of the shore crab (Carcinus maenas L.). To determine the source of this variation (genetic or environmental), morphological and genetic data were analysed from crabs collected from eight sites around the coast of the UK. Ten morphometric traits were measured from over 800 crabs and the degree of morphological similarity among sites was calculated using multivariate techniques. Allozyme electrophoresis was used to investigate patterns of genetic similarity. Extensive morphological variability was detected: eight out of the ten morphometric traits analysed were useful when discriminating between crabs from each site. Discriminant function analysis revealed that over 35% of individuals could be classified to their site of origin on the basis of their morphology. In contrast, the allozyme analysis revealed low levels of genetic variability, both within the meta-population and among the crab population at each site. Pairwise comparisons revealed a moderate correlation between the degree of morphological and genetic similarity of crabs at each site, which suggests that the observed phenotypic variability has a genetic component. However, only around 20% of the phenotypic variability detected was associated with the patterns of genetic similarity. This means that patterns of morphological variability in this species are largely determined by the local environmental conditions: local factors could have a within-generation selective influence on mean trait values or C. maenas may exhibit phenotypic plasticity.     

灰木莲种源幼苗叶片性状表型多样性分析

以越南收集的12个灰木莲（Manglietia conifera Dandy）种源为材料,对叶片形态性状（叶长、叶宽、叶面积、叶周长、长宽比、叶柄）以及微形态特征（气孔器密度、气孔器长、气孔器宽、气孔器面积、长宽比）进行测定,采用方差分析、变异系数、相关分析和主成分分析等方法进行分析。结果表明：灰木莲不同种源间的叶表型性状存在显著差异;种源中,LC2的叶形态性状的平均变异系数最大（22.09%）,TQ2的变异系数最小（12.76%）;叶表型性状中,叶面积的变异系数最大（28.60%）,气孔器宽的变异系数最小（5.19%）;相关性分析结果表明叶表型性状间存在显著或极显著的相关关系,而地理因素中经度与叶周长显著相关,经纬度与叶长宽比呈显著相关,海拔与叶表型性状间的相关性不显著（P<0.05）;主成分分析表明前3个主成分的累计贡献率达到了89.29%,基本代表原始性状的全部信息。灰木莲12个种源经聚类分析在欧式距离5阈值处可分为4类。灰木莲种源间的叶表型性状存在着丰富的变异,纬度对灰木莲叶形态特征有明显影响,灰木莲种源间和性状间的变异程度存在着差异,本研究为灰木莲遗传改良提供理论依据。     

Embryological aspects of inducible morphological defenses in Daphnia

Many cases of predator-induced morphological plasticity in daphnids are well studied examples of inducible defenses. However, little is known about the early development of these sometimes conspicuous traits. We compared for the first time in five different Daphnia species the embryonic development of predator-induced and noninduced animals using scanning electron microscopy (SEM). We observed significant morphological changes in the last embryonic stage in helmet formation in Daphnia cucullata and in neck-pedestal development in Daphnia pulex. In contrast, no morphological changes could be found during embryogenesis between induced and noninduced Daphnia lumholtzi, D. longicephala, and D. ambigua. Strategies for initiating the defensive traits differ among Daphnia species because of trade-offs between environmental requirements and developmental constraints. Some general features of Daphnia embryonic development are described using SEM. All Daphnia embryos have to shed at least three different membranes before leaving the brood pouch of the mother. After the embryos shed the third membrane, chemosensillae that are likely able to detect predator-released chemicals are exposed to the olfactory environment.     

Primitive versus derived traits in the developmental program of the vertebrate head: views from cyclostome developmental studies

Evolution can be viewed as a series of changes in the developmental program along the phylogenetic tree. To better understand the early evolution of the vertebrate skull, we can use the embryos of the cyclostome species as models. By comparing the cyclostome developmental patterns with those of gnathostomes, it becomes possible to distinguish the primitive and derived parts of the developmental program as taxon-specific traits. These traits are often recognizable as developmental constraints that define taxa by biasing the developmental trajectories within a certain limited range, resulting in morphological homologies in adults. These developmental constraints are distributed on the phylogenetic tree like the morphological character states of adult animals and are associated with specific regions of the tree. From this perspective, we emphasize the importance of considering gene expression and embryonic anatomy as the mechanistic bases that can result in homologous or nonhomologous morphological patterns at later developmental stages. Taking the acquisition of the jaw and trabecula cranii as examples, we demonstrate that a set of embryonic features can be coupled or decoupled during evolution and development. When they are coupled, they exert an ancestral developmental constraint that results in homologous morphological patterns, and when they are decoupled, the ancestral constraints tend to be abandoned, generating a new body plan. The heterotopy behind the specification of the oral domain is an example of decoupling, based on shifted tissue interactions. We also stress the importance of "developmental burden" in determining the sequential order of changes through evolution.     

Protamine withdrawal from human sperm nuclei following heterologous ICSI into hamster oocytes

During late stages of spermatogenesis in mammals, most histones bound to DNA are replaced by protamines (PRM), which results in formation of supercondensed and genetically inert sperm chromatin. At fertilization, mature spermatozoon penetrates oocyte and chromatin is remodeled "back" from nucleoprotamine to nucleohistone state. While being crucial for activation of male genome and ultimately for initiation of embryonic development, this process is poorly studied, especially in humans. Data on model animals concerning PRM to histones exchange post fertilization are few and contradictory. As direct experimentation with human embryos is impossible due to ethical, legal and technical reasons, we evaluate the timing and mode of PRM removal in a heterologous ICSI system using hamster ova injected with human sperm. Localization of human PRM 1 and 2 in hybrid zygotes was established using immunofluorescence. We observed a marked zygote to zygote variability in male pronuclei size for any time point post ICSI and demonstrated that PRM removal correlates with the developing pronuclei area rather than time after injection. Overall, the disappearance of protamines from sperm is rather rapid and most likely completed within 1 hr. We propose that the critical characteristic influencing PRM removal after heterologous fertilization is the intrinsic heterogeneity of the human sperm population. The same yet unexplored variance may be one of the reasons for canceled, delayed or aberrant early embryonic development during natural or artificial fertilization in humans.     

Testing evolutionary hypotheses about the phylotypic period of zebrafish

Vertebrate embryos pass through a period of morphological similarity, the phylotypic period. Since Haeckel's biogenetic law of recapitulation, proximate and ultimate evolutionary causes of such similarity of embryos were discussed. We test predictions about changes in phenotypic and genetic variances that were derived from three hypotheses about the evolutionary origin of the phylotypic stage, i.e. random, epigenetic effects, and stabilizing selection. The random hypothesis predicts increasing values for phenotypic variances and stable or increasing values for genetic variances; the epigenetic effects hypothesis predicts declining values for phenotypic variances but stable or increasing values of genetic variances, and the stabilizing selection predicts stable phenotypic variances but decreasing genetic variances. We studied zebrafish as a model species, because it can be bred in large numbers as necessary for a quantitative genetics breeding design. A half-sib breeding scheme provided estimates of additive genetic variances from 11 embryonic characters from 12 through to 24 hr after fertilization, i.e. before, during (15-19 hr), and after the phylotypic period. Because additive genetic variances are size dependent, we calculated narrow-sense heritabilities as a size independent gauge of genetic contributions to the phenotype. The results show declining phenotypic variances and stable heritabilities. In conclusion, we reject the random and the stabilizing selection hypotheses and favor ideas about epigenetic effects that constrain the early embryonic development. Additive genetic variance during the phylotypic stage makes it accessible for evolution, thus explaining in a simple and straightforward way why the phylotypic period differs among vertebrates in timing, duration, and morphologies.     

皂荚南方天然群体种实表型多样性

以系统揭示表型变异程度和变异规律为目的, 对皂荚(Gleditsia sinensis)南方分布区的10个天然群体的11个种实性状进行了比较分析。采用方差分析、多重比较、相关分析等多种分析方法, 对群体间和群体内的表型多样性以及与地理、环境因子的相关性进行了讨论。方差分析结果表明; 皂荚果实、种子等性状在群体间和群体内存在丰富的变异, 11个性状在群体间、群体内均达显著差异; 荚果性状在群体间和群体内的变异均大于种子性状, 11个性状的平均表型分化系数为20.42%, 群体内的变异(32.28%)大于群体间的变异(7.19%), 群体内的变异是皂荚的主要变异来源; 皂荚各性状平均变异系数为11.20%, 变异幅度为4.55%-18.38%。群体间荚果的变异(14.75%)高于群体间种子变异(6.95%), 表明种子变异稳定性高。荚果和种子各性状之间多呈极显著或显著正相关, 表现为荚果越大, 则种子越大, 种子的千粒重也越大; 荚果表现为同地理经度的南北变异, 种子则表现为同地理纬度的东西变异。研究结果为进一步开展皂荚遗传育种、保护生物学研究和皂荚种质资源利用奠定了基础。     

Linking stream ecology with morphological variability in a native freshwater fish from semi‐arid Australia

Environmental variation is a potent force affecting phenotypic expression. While freshwater fishes have provided a compelling example of the link between the environment and phenotypic diversity, few studies have been conducted with arid‐zone fishes, particularly those that occur in geographically isolated regions where species typically inhabit intermittent and ephemeral creeks. We investigated morphological variation of a freshwater fish (the western rainbowfish, Melanotaenia australis) inhabiting creeks in the Pilbara region of northwest Australia to determine whether body shape variation correlated with local environmental characteristics, including water velocity, habitat complexity, predator presence, and food availability. We expected that the geographic isolation of creeks within this arid region would result in habitat‐specific morphological specializations. We used landmark‐based geometric morphometrics to quantify the level of morphological variability in fish captured from 14 locations within three distinct subcatchments of a major river system. Western rainbowfish exhibited a range of morphologies, with variation in body depth accounting for a significant proportion (>42%) of the total variance in shape. Sexual dimorphism was also apparent, with males displaying deeper bodies than females. While the measured local habitat characteristics explained little of the observed morphological variation, fish displayed significant morphological differentiation at the level of the subcatchment. Local adaptation may partly explain the geographic patterns of body shape variation, but fine‐scale genetic studies are required to disentangle the effects of genetic differentiation from environmentally determined phenotypic plasticity in body shape. Developing a better understanding of environment–phenotype relationships in species from arid regions will provide important insights into ecological and evolutionary processes in these unique and understudied habitats.     

Patterns of phenotypic covariation and correlation in modern humans as viewed from morphological integration

Proportionality of phenotypic and genetic distance is of crucial importance to adequately focus on population history and structure, and it depends on the proportionality of genetic and phenotypic covariance. Constancy of phenotypic covariances is unlikely without constancy of genetic covariation if the latter is a substantial component of the former. If phenotypic patterns are found to be relatively stable, the most probable explanation is that genetic covariance matrices are also stable. Factors like morphological integration account for such stability. Morphological integration can be studied by analyzing the relationships among morphological traits. We present here a comparison of phenotypic correlation and covariance structure among worldwide human populations. Correlation and covariance matrices between 47 cranial traits were obtained for 28 populations, and compared with design matrices representing functional and developmental constraints. Among-population differences in patterns of correlation and covariation were tested for association with matrices of genetic distances (obtained after an examination of 10 Alu-insertions) and with Mahalanobis distances (computed after craniometrical traits). All matrix correlations were estimated by means of Mantel tests. Results indicate that correlation and covariance structure in our species is stable, and that among-group correlation/covariance similarity is not related to genetic or phenotypic distance. Conversely, genetic and morphological distance matrices were highly correlated. Correlation and covariation patterns were largely associated with functional and developmental factors, which probably account for the stability of covariance patterns.     

Phenotypic variation in smooth softshell turtles (<Emphasis Type="Italic">Apalone mutica</Emphasis>) from eggs incubated in constant versus fluctuating temperatures

Temperatures experienced during embryonic development elicit well-documented phenotypic variation in embryonic and neonatal animals. Most research, however, has only considered the effects of constant temperatures, even though developmental temperatures in natural settings fluctuate considerably on a daily and seasonal basis. A laboratory study of 15 clutches of smooth softshell turtles (Apalone mutica) was conducted to explicitly examine the influence of thermal variance on phenotypic variation. Holding mean temperature constant and eliminating substrate moisture effects permitted a clear assessment of the impact of thermal variance on hatching success, incubation length, hatchling body size, swimming speed, and righting time. Incubation length and swimming speed varied significantly among temperature treatments. Both traits tended to increase with increasing thermal variance during embryonic development. Clutch significantly affected all traits examined, except righting time, even after accounting for the effects of initial egg mass. These results highlight the importance of accounting for the impact of both thermal mean and variance on phenotypic variation. The findings also strengthen the increasing recognition of maternal clutch effects as critical factors influencing phenotypic variation in neonatal animals.     

Characterization of NF-kappa B/I kappa B proteins in zebra fish and their involvement in notochord development

Although largely involved in innate and adaptive immunity, NF-kappa B plays an important role in vertebrate development. In chicks, the inactivation of the NF-kappa B pathway induces functional alterations of the apical ectodermal ridge, which mediates limb outgrowth. In mice, the complete absence of NF-kappa B activity leads to prenatal death and neural tube defects. Here, we report the cloning and characterization of NF-kappa B/I kappa B proteins in zebra fish. Despite being ubiquitously expressed among the embryonic tissues, NF-kappa B/I kappa B members present distinct patterns of gene expression during the early zebra fish development. Biochemical assays indicate that zebra fish NF-kappa B proteins are able to bind consensus DNA-binding (kappa B) sites and inhibitory I kappa B alpha proteins from mammals. We show that zebra fish I kappa B alphas are degraded in a time-dependent manner after induction of transduced murine embryo fibroblasts (MEFs) and that these proteins are able to rescue NF-kappa B activity in I kappa B alpha(-/-) MEFs. Expression of a dominant-negative form of the murine I kappa B alpha (mI kappa B alpha M), which is able to block NF-kappa B in zebra fish cells, interferes with the notochord differentiation, generating no tail (ntl)-like embryos. This phenotype can be rescued by coinjection of the T-box gene ntl (Brachyury homologue), which is typically required for the formation of posterior mesoderm and axial development, suggesting that ntl lies downstream of NF-kappa B . We further show that ntl and Brachyury promoter regions contain functional kappa B sites and NF-kappa B can directly modulate ntl expression. Our study illustrates the conservation and compatibility of NF-kappa B/I kappa B proteins among vertebrates and the importance of NF-kappa B pathway in mesoderm formation during early embryogenesis.