Ecological constraints on diversification in a model adaptive radiation

Taxonomic diversification commonly occurs through adaptive radiation, the rapid evolution of a single lineage into a range of genotypes or species each adapted to a different ecological niche. Radiation size (measured as the number of new types) varies widely between phylogenetically distinct taxa and between replicate radiations within a single taxon where the ecological opportunities available seem to be identical. Here we show how variation in energy input (productivity) and environmental disturbance combine to determine the extent of diversification in a single radiating lineage of Pseudomonas fluorescens adapting to laboratory conditions. Diversity peaked at intermediate rates of both productivity and disturbance and declined towards the extremes in a manner reminiscent of well-known ecological patterns. The mechanism responsible for the decrease in diversity arises from pleiotropic fitness costs associated with niche specialization, the effects of which are modulated by gradients of productivity and disturbance. Our results indicate that ecological gradients may constrain the size of adaptive radiations, even in the presence of the strong diversifying selection associated with ecological opportunity, by decoupling evolutionary diversification from ecological coexistence.     

Ecological opportunity and sexual selection together predict adaptive radiation

A fundamental challenge to our understanding of biodiversity is to explain why some groups of species undergo adaptive radiations, diversifying extensively into many and varied species, whereas others do not. Both extrinsic environmental factors (for example, resource availability, climate) and intrinsic lineage-specific traits (for example, behavioural or morphological traits, genetic architecture) influence diversification, but few studies have addressed how such factors interact. Radiations of cichlid fishes in the African Great Lakes provide some of the most dramatic cases of species diversification. However, most cichlid lineages in African lakes have not undergone adaptive radiations. Here we compile data on cichlid colonization and diversification in 46 African lakes, along with lake environmental features and information about the traits of colonizing cichlid lineages, to investigate why adaptive radiation does and does not occur. We find that extrinsic environmental factors related to ecological opportunity and intrinsic lineage-specific traits related to sexual selection both strongly influence whether cichlids radiate. Cichlids are more likely to radiate in deep lakes, in regions with more incident solar radiation and in lakes where there has been more time for diversification. Weak or negative associations between diversification and lake surface area indicate that cichlid speciation is not constrained by area, in contrast to diversification in many terrestrial taxa. Among the suite of intrinsic traits that we investigate, sexual dichromatism, a surrogate for the intensity of sexual selection, is consistently positively associated with diversification. Thus, for cichlids, it is the coincidence between ecological opportunity and sexual selection that best predicts whether adaptive radiation will occur. These findings suggest that adaptive radiation is predictable, but only when species traits and environmental factors are jointly considered.     

Climate-driven population divergence in sex-determining systems

Sex determination is a fundamental biological process, yet its mechanisms are remarkably diverse. In vertebrates, sex can be determined by inherited genetic factors or by the temperature experienced during embryonic development. However, the evolutionary causes of this diversity remain unknown. Here we show that live-bearing lizards at different climatic extremes of the species' distribution differ in their sex-determining mechanisms, with temperature-dependent sex determination in lowlands and genotypic sex determination in highlands. A theoretical model parameterized with field data accurately predicts this divergence in sex-determining systems and the consequence thereof for variation in cohort sex ratios among years. Furthermore, we show that divergent natural selection on sex determination across altitudes is caused by climatic effects on lizard life history and variation in the magnitude of between-year temperature fluctuations. Our results establish an adaptive explanation for intra-specific divergence in sex-determining systems driven by phenotypic plasticity and ecological selection, thereby providing a unifying framework for integrating the developmental, ecological and evolutionary basis for variation in vertebrate sex determination.     

Nuclear DNA sequences detect species limits in ancient moa

Ancient DNA studies have typically used multi-copy mitochondrial DNA sequences. This is largely because single-locus nuclear genes have been difficult to recover from sub-fossil material, restricting the scope of ancient DNA research. Here, we have isolated single-locus nuclear DNA markers to assign the sex of 115 extinct moa and, in combination with a mitochondrial DNA phylogeny, tested competing hypotheses about the specific status of moa taxa. Moa were large ratite birds that showed extreme size variation both within and among species. For some taxa, this large variation was hypothesized to represent sexual dimorphism, while for others it was argued to reflect the existence of different species. Our results show that moa were characterized by extreme reverse sexual dimorphism and as a result we have been able to clarify the number of moa species. For example, we show that the three recognized 'species' of Dinornis comprised only two monophyletic groups and that two of these 'species' comprised individuals of one sex only. This study also illustrates that single-locus nuclear DNA sequences can be consistently recovered from ancient material.     

The ecological cost of sex

Why sex prevails in nature remains one of the great puzzles of evolution. Sexual reproduction has an immediate cost relative to asexual reproduction, as males only express their contribution to population growth through females. With no males to sustain, an asexual mutant can double its relative representation in the population in successive generations. This is the widely accepted 'twofold cost of males'. Many studies have attempted to explain how sex can recoup this cost from fitness benefits associated with the recombination of parental genotypes, but these require complex biological environments that cycle over evolutionary timescales. In contrast, we have considered the ecological dynamics that govern asexual invasion. Here we show the existence of a threshold growth rate for the sexual population, above which the invasion is halted by intraspecific competition. The asexual population then exerts a weaker inhibitory effect on the carrying capacity of the sexual population than on its own carrying capacity. The stable outcome of this is coexistence on a depleted resource base. Under these ecological circumstances, longer-term benefits of sex may eventually drive out the asexual competitor.     

太行山猕猴牙齿性差的分布模式和相关性研究

为了解太行山猕猴牙齿性差程度、分布区域和模式,研究各个牙齿之间的相关程度,研究犬齿及邻近牙齿的相关性.选41例太行山猕猴(雌29,雄12)牙齿标本.观测变量为牙齿近中-远中径(MD)和颊-舌径(BL;)数据经SPSS12.0统计软件处理.相关分析采用双变量简单相关和偏相关分析.结果发现:太行山猕猴牙齿性差主要表现在犬齿"性差区"和第3臼齿.靠近犬齿的牙齿其性差较大.偏相关研究结果表明,犬齿对其邻近的牙齿的影响较小,形态上相近的牙齿具有更大的相关性.牙齿性差大小主要受遗传等因素控制.     

五种蝰科蛇尾长的两性异形和种间差异

测定五种分布于浙江的蝰科蛇类的体长和尾长,检测尾长的两性异形和种间差异。五种蝰科蛇的两性个体尾长均与体长呈显著的正相关,五种蝰科蛇间体长差异显著。协方差分析显示雄性的尾长显著大于雌性。除烙铁头、竹叶青之间其余相对尾长的差异显著。     

猕猴骶骨性别判别分析

目的:探讨太行山猕猴骶骨指数性差.材料和方法:成年猕猴骶骨标本55例(雄17,雌38).选择骶骨5个线性变量和4个指数变量;统计处理采用ANOVA、多元判别分析和二元逻辑回归分析.结果:大部分骶骨线性变量存在性差(P0.05),雄性大于雌性;猕猴骶骨相对宽指数性差显著(P0.01),雌性大于雄性.多元判别分析性别正确判别率为78.2%~87.3%.逻辑回归分析性别正确判别率为83.6%~87.3%.结论:骶骨变量性差显著,骶骨体长和骶骨相对宽性别判别的能力较强.     

MHC II DRB variation and trans-species polymorphism in the golden snub-nosed monkey (Rhinopithecus roxellana)

Genetic variation is generally believed to be important in studying endangered species’ adaptive potential.Early studies assessed genetic diversity using nearly neutral markers,such as microsatellite loci and mitochondrial DNA(mtDNA),which are very informative for phylogenetic and phylogeographic reconstructions.However,the variation at these loci cannot provide direct information on selective processes involving the interaction of individuals with their environment,or on the capability to resist continuously evolving pathogens and parasites.The importance of genetic diversity at informative adaptive markers,such as major histocompatibility complex(MHC) genes,is increasingly being realized,especially in endangered,isolated species.Small population size and isolation make the golden snub-nosed monkey(Rhinopithecus roxellana) particularly susceptible to genetic variation losses through inbreeding and restricted gene flow.In this study,we compared the genetic variation and population structure of microsatellites,mtDNA,and the most relevant adaptive region of the MHC II-DRB genes in the golden snub-nosed monkey.We examined three Chinese R.roxellana populations and found the same variation patterns in all gene regions,with the population from Shennongjia population,Hubei Province,showing the lowest polymorphism among three populations.Genetic drift that outweighed balancing selection and the founder effect in these populations may explain the similar genetic variation pattern found in these neutral and adaptive genes.     

六种游蛇属蛇尾长的两性异形和种间差异

测定六种分布于浙江的游蛇属蛇类的体长和尾长,检测尾长的两性异形和种间差异.六种游蛇属蛇的两性个体尾长均与体长呈显著的正相关,六种游蛇属蛇间体长差异显著.协方差分析显示渔游蛇、草游蛇、虎斑游蛇的雄性的尾长显著大于雌性,水赤链游蛇、锈链游蛇、乌游蛇的尾长无显著的两性差异.种间相对尾长的差异显著.     

Sexual selection enables long-term coexistence despite ecological equivalence

Empirical data indicate that sexual preferences are critical for maintaining species boundaries, yet theoretical work has suggested that, on their own, they can have only a minimal role in maintaining biodiversity. This is because long-term coexistence within overlapping ranges is thought to be unlikely in the absence of ecological differentiation. Here we challenge this widely held view by generalizing a standard model of sexual selection to include two ubiquitous features of populations with sexual selection: spatial variation in local carrying capacity, and mate-search costs in females. We show that, when these two features are combined, sexual preferences can single-handedly maintain coexistence, even when spatial variation in local carrying capacity is so slight that it might go unnoticed empirically. This theoretical study demonstrates that sexual selection alone can promote the long-term coexistence of ecologically equivalent species with overlapping ranges, and it thus provides a novel explanation for the maintenance of species diversity.     

草地植物功能性状变异的程度和来源

分析了内蒙古草地植物的6个功能性状在种间、种内不同种群间和同一种群内不同个体间的变异.结果表明,除株高外其余5个功能性状的变异主要来自种间,但种内变异占总体变异的比例不能被忽视.不同功能性状的种内和种间变异的程度和来源不同,说明不同物种、不同性状对环境变化的响应不同.因此利用功能性状进行各种研究时,应该充分考虑种内变异的影响.     

Fruitless specifies sexually dimorphic neural circuitry in the Drosophila brain

The Drosophila fruitless (fru) gene product Fru has been postulated to be a neural sex determination factor that directs development of the central nervous system (CNS), thereby producing male-typical courtship behaviour and inducing male-specific muscle. Male-specific Fru protein is expressed in small groups of neurons scattered throughout the CNS of male, but not female, Drosophila. Collectively, these observations suggest that Fru 'masculinizes' certain neurons, thereby establishing neural substrates for male-typical behaviour. However, specific differences between neurons resulting from the presence or absence of Fru are unknown. Previous studies have suggested that Fru might result in sexual differences in the CNS at the functional level, as no overt sexual dimorphism in CNS structure was discernible. Here we identify a subset of fru-expressing interneurons in the brain that show marked sexual dimorphism in their number and projection pattern. We also demonstrate that Fru supports the development of neurons with male-specific dendritic fields, which are programmed to die during female development as a result of the absence of Fru. Thus, Fru expression can produce a male-specific neural circuit, probably used during heterosexual courtship, by preventing cell death in identifiable neurons.     

Intensity of sexual selection along the anisogamy-isogamy continuum

Research into the evolution of giant sperm has uncovered a paradox within the foundations of sexual selection theory. Postcopulatory sexual selection on males (that is, sperm competition and cryptic female choice) can lead to decreased sperm numbers by favouring the production of larger sperm. However, a decline in sperm numbers is predicted to weaken selection on males and increase selection on females. As isogamy is approached (that is, as investment per gamete by males approaches that by females), sperm become less abundant, ova become relatively less rare, and competition between males for fertilization success is predicted to weaken. Sexual selection for longer sperm, therefore, is expected to be self limiting. Here we examine this paradox in Drosophila along the anisogamy-isogamy continuum using intraspecific experimental evolution techniques and interspecific comparative techniques. Our results confirm the big-sperm paradox by showing that the sex difference in sexual selection gradients decreases as sperm size increases. However, a resolution to the paradox is provided when this finding is interpreted in concert with the 'opportunity for selection' and the 'opportunity for sexual selection'. Furthermore, we show that most of the variation in measures of selection intensity is explained by sperm length and relative investment in sperm production.     

基于能值理论的自然生态系统经济价值研究——以大巴山南坡雪宝山自然生态系统为例

以能值理论为基础,以大巴山南坡雪宝山自然生态系统为研究对象,对自然生态系统的经济价值进行了研究.研究结果表明,雪宝山自然生态系统的经济价值约为2.64×107US＄/a,其中物质产品和生态服务(不含生物多样性)的经济价值分别为2.26×107US＄/a、3.67×106US＄/a,生物多样性的经济价值约为7.75×109US＄.这说明在评估自然生态系统的经济价值时,生态服务和生物多样性的经济价值不容忽视.     

An extant cichlid fish radiation emerged in an extinct Pleistocene lake

The haplochromine cichlid fish of the East African Great Lakes represent some of the fastest and most species-rich adaptive radiations known, but rivers in most of Africa accommodate only a few morphologically similar species of haplochromine cichlid fish. This has been explained by the wealth of ecological opportunity in large lakes compared with rivers. It is therefore surprising that the rivers of southern Africa harbour many, ecologically diverse haplochromines. Here we present genetic, morphological and biogeographical evidence suggesting that these riverine cichlids are products of a recent adaptive radiation in a large lake that dried up in the Holocene. Haplochromine species richness peaks steeply in an area for which geological data reveal the historical existence of Lake palaeo-Makgadikgadi. The centre of this extinct lake is now a saltpan north of the Kalahari Desert, but it once hosted a rapidly evolving fish species radiation, comparable in morphological diversity to that in the extant African Great Lakes. Importantly, this lake seeded all major river systems of southern Africa with ecologically diverse cichlids. This discovery reveals how local evolutionary processes operating during a short window of ecological opportunity can have a major and lasting effect on biodiversity on a continental scale.     

Sex determination compared in Drosophila and Caenorhabditis

Fruitflies and nematodes show many similarities in the general organization of the gene networks that control sexual dimorphism and dosage compensation. In contrast, the underlying molecular mechanisms appear to be very different in these two species. Developmental processes such as sex determination need not be strongly conserved in evolution.     

Large-scale processes and the Asian bias in species diversity of temperate plants

An important issue in the study of biodiversity is the extent to which global patterns of species richness reflect large-scale processes and historical contingencies. Ecological interactions in local assemblages may constrain the number of species that can coexist, but differences in diversity in similar habitats within different regions (diversity anomalies) suggest that this limit is not firm. Variation in rate of species production could influence regional and perhaps local diversity independently of the ecological capacity of an area to support coexisting species, thereby creating diversity anomalies. Temperate Zone genera of plants that are disjunct between similar environments in eastern Asia and eastern North America (EAS-ENA) have twice as many species in Asia as in North America. Because lineages of these genera in Asia and North America are mostly sister pairs, they share a common history of adaptation and ecological relationship before disjunction. Thus, the diversity anomaly in EAS-ENA genera is not an artefact of taxon or habitat sampling but reflects differences in the net diversification (speciation-extinction) of the lineages in each of the continents. Here we propose that the most probable cause of the EAS-ENA anomaly in diversity is the extreme physiographical heterogeneity of temperate eastern Asia, especially compared with eastern North America, which in conjunction with climate and sea-level change has provided abundant opportunities for evolutionary radiation through allopatric speciation.     

Multiple routes to mammalian diversity

The radiation of the mammals provides a 165-million-year test case for evolutionary theories of how species occupy and then fill ecological niches. It is widely assumed that species often diverge rapidly early in their evolution, and that this is followed by a longer, drawn-out period of slower evolutionary fine-tuning as natural selection fits organisms into an increasingly occupied niche space. But recent studies have hinted that the process may not be so simple. Here we apply statistical methods that automatically detect temporal shifts in the rate of evolution through time to a comprehensive mammalian phylogeny and data set of body sizes of 3,185 extant species. Unexpectedly, the majority of mammal species, including two of the most speciose orders (Rodentia and Chiroptera), have no history of substantial and sustained increases in the rates of evolution. Instead, a subset of the mammals has experienced an explosive increase (between 10- and 52-fold) in the rate of evolution along the single branch leading to the common ancestor of their monophyletic group (for example Chiroptera), followed by a quick return to lower or background levels. The remaining species are a taxonomically diverse assemblage showing a significant, sustained increase or decrease in their rates of evolution. These results necessarily decouple morphological diversification from speciation and suggest that the processes that give rise to the morphological diversity of a class of animals are far more free to vary than previously considered. Niches do not seem to fill up, and diversity seems to arise whenever, wherever and at whatever rate it is advantageous.     

体育在青少年健康性心理形成中的应用研究

本研究通过体育的干预来促进青少年健康性心理的形成,运用文献资料、实验和问卷调查统计等方法研究青少年在参加不同体育项目练习后性心理的变化情况。研究表明,不同的体育项目对于青少年性度的养成、性体像的重塑和性角色的认同具有明显的促进作用,对于增强学生的自信心和提高性心理的健康水平有着非常重要意义,同时也为性教育的普及提供了参考依据。