Evolutionary trade-offs between reproduction and dispersal in populations at expanding range boundaries

During recent climate warming, some species have expanded their ranges northwards to keep track of climate changes. Evolutionary changes in dispersal have been demonstrated in these expanding populations and here we show that increased dispersal is associated with reduced investment in reproduction in populations of the speckled wood butterfly, Pararge aegeria. Evolutionary changes in flight versus reproduction will affect the pattern and rate of expansion at range boundaries in the future, and understanding these responses will be crucial for predicting the distribution of species in the future as climates continue to warm.     

Evolutionarily stable allocation to vegetative and sexual reproduction in plants

I apply the resource allocation approach to construct an ESS model aimed at predicting the rate of vegetative and sexual reproduction in plants. The model provides an integrated explanation of why the mode of reproduction should vary between populations and species, and what factors should affect the trade-off between ramet and seed production. It follows that seed production, usually considered to be less effective and more costly than ramet production, may be maintained in the population. The production of ramets should be favored when their recruitment rate is high or the costs are low. The intensity of vegetative reproduction is also related to conditions which affect seed production directly. The rate of ramet production should be inversely correlated with factors favoring sexual reproduction, such as lower cost of seeds or high opportunity for colonization of new patches, etc. Under such conditions it is beneficial to allocate more resources available for reproduction into seeds, at the expense of ramet production. The model emphasizes the need to consider both modes of plant reproduction whenever one of them is theoretically or empirically studied.     

Sex-Specific Functional Specialization and the Evolutionary Rates of Essential Fertility Genes

Genes related to sex and reproduction are known to evolve rapidly, however, the mechanism for rapid evolutionary change is proving to be more complex than a simple relaxation of selective constraint. We compared the divergence between orthologous human and mouse fertility genes according to their degree of dispensability as suggested by mouse knockout mutation phenotypes. The dataset consisted of 161 orthologous genes affecting fertility and 803 orthologous genes affecting viability. We find that essential fertility genes affecting both sexes evolve at a similar rate as essential viability genes, but that within sexes the degree of dispensability is not an important factor affecting the rate of fertility gene evolution. We also find no difference in the evolutionary rates of fertility genes that affect the male versus the female, however, there are a greater number of sterility genes that affect the male. Generally there are a significantly greater number of fertility genes that affect one sex rather than both, suggesting that fertility genes tend toward sex-specific functions, particularly in the male. Our findings support the hypothesis that the rapid evolution of sex- and reproduction-related genes is facilitated through an increased specialization of gene function and that dispensability is not a major factor determining their evolutionary rate. Electronic Supplementary Material Electronic Supplementary material is available for this article at and accessible for authorised users. [Reviewing Editor: Dr. Willie J. Swanson]     

Combined analysis of Perca fluviatilis reproductive performance and oocyte proteomic profile

The success of reproduction depends greatly upon gamete quality, especially oocytes which carry most of the molecular material necessary for early embryogenesis. However, it remains difficult to find relevant morphologic and/or biochemical parameters to assess oocyte quality and thus have a reliable prediction of the reproduction performance. To understand which criteria are the most reliable to assess the reproductive success of the Eurasian perch (Perca fluviatilis), we measured 14 parameters characterizing female, spawn, oocyte, and embryonic or larval development on 20 independent spawn. A data analysis allowed the definition of two clusters of spawn with different larval characteristics: the first cluster was composed of spawn which led mainly to strong large larvae presenting a low deformity rate, while the second cluster rather corresponds to spawn leading to smaller and weaker larvae with a higher deformity rate. Moreover, a third cluster (unfertilized spawn) was studied. Our analysis revealed that most of the prefertilization biological traits that we studied appeared poorly relevant to predict larval features, proper embryonic development and deformity occurrences. We thus performed a large scale proteomic analysis to highlight proteins differently expressed in each spawn cluster. A 2D-DIGE study followed by an MS/MS spectrometry allowed the identification of 32 proteins involved in several biological functions and differently expressed between spawn clusters. Among them, proteins involved in cell response to the oxidative stress, as well as energetic metabolism, heat shock proteins and Vitellogenins are of particular interest. Several functions appear specific to a spawn cluster and could thus explain their corresponding reproduction performance. In the future, proteins involved in those cellular mechanisms may constitute molecular markers predictive of the reproduction performance in Perca fluviatilis.     

Mutation,selection, and ancestry in branching models: a variational approach

We consider the evolution of populations under the joint action of mutation and differential reproduction, or selection. The population is modelled as a finite-type Markov branching process in continuous time, and the associated genealogical tree is viewed both in the forward and the backward direction of time. The stationary type distribution of the reversed process, the so-called ancestral distribution, turns out as a key for the study of mutation–selection balance. This balance can be expressed in the form of a variational principle that quantifies the respective roles of reproduction and mutation for any possible type distribution. It shows that the mean growth rate of the population results from a competition for a maximal long-term growth rate, as given by the difference between the current mean reproduction rate, and an asymptotic decay rate related to the mutation process; this tradeoff is won by the ancestral distribution. We then focus on the case when the type is determined by a sequence of letters (like nucleotides or matches/mismatches relative to a reference sequence), and we ask how much of the above competition can still be seen by observing only the letter composition (as given by the frequencies of the various letters within the sequence). If mutation and reproduction rates can be approximated in a smooth way, the fitness of letter compositions resulting from the interplay of reproduction and mutation is determined in the limit as the number of sequence sites tends to infinity. Our main application is the quasispecies model of sequence evolution with mutation coupled to reproduction but independent across sites, and a fitness function that is invariant under permutation of sites. In this model, the fitness of letter compositions is worked out explicitly. In certain cases, their competition leads to a phase transition.      

Evolution of senescence: Natural increase of populations displaying gompertz- or power-law death rates and constant or age-dependent maternity rates

Several hypothetical populations which differ in degrees of senescence are compared with respect to their rates of natural increase. The rate of natural increase is employed as a measure of selective advantage. The populations are characterized by their maternity and death rates, expressed as functions of age. Maternity rates are described by constant or quasi-human, age-dependent functions. Death rates are described by constant, Gompertzian (exponential) or power functions. Longevity functions, representing the probability of survival to a specific age, are obtained by integrating the death rate functions. The degree of senescence of a population is measured by the rapidity of ascent of its death-rate function or by the rectangularity of its longevity function. The increase in death rate late in life which constitutes senescence is compensated by a decrease in death rate early in life. The balance between the two changes in rate is, by assumption, such that the mean value of the longevity function is independent of the degree of senescence. This assumption makes it possible to separate the effects produced by the evolution of senescence from those caused by changes in longevity.The rate of natural increase is obtained by numerical solution of an integral characteristic equation. The results show that senescence is advantageous in all populations except those in which the maternity function is constant and the size is declining at a rapid rate. When the parameters entering into the longevity functions have values such that the functions approximate human longevity data, the improvement in the rate of natural increase resulting from senescence closely approaches limiting values obtained with the use of a precisely rectangular longevity function. Other results support the observation that reproduction at an early age confers greater selective advantage than equivalent reproduction later in life.     

单宁酸对布氏田鼠能量代谢的影响

为了解单宁酸对成年布氏田鼠(Lasiopodomys bandtii**)能量代谢和产热的影响,本文采用含0、3.3%和6.6%单宁酸浓度的食物饲喂布氏田鼠21 d,对其体重、基础代谢率、非颤抖性产热和能量收支等进行了测定。代谢率采用封闭式流体压力呼吸计测定;非颤抖性产热用皮下注射去甲肾上腺素诱导;能量摄人采用食物平衡法测定。结果发现:(1) 单宁酸食物对布氏田鼠的体重没有明显影响;(2)取食含6.6%单宁酸食物的动物的基础代谢率于第10 d高于对照组。20 d时,3组动物的基础代谢率没有显著差异;(3) 单宁酸食物对非颤抖性产热没有显著影响;(4) 食用含单宁酸食物的动物的摄人能和消化能于第10 d显著低于对照组,但第20 d时则差异不显著。这些结果表明:布氏田鼠的基础代谢率和能量摄入对单宁酸的反应具有时段性,短期内能量消耗增加,随着动物对食物的适应,生理功能恢复到正常水平。     

Adaptive advantages of patterns of growth and asexual reproduction of the sea anemone Metridium senile (L.) in intertidal and submerged populations

Individual size, rate of growth, and mode and frequency of asexual reproduction are life-history traits of primary importance for sea anemones. These traits determine sexual reproductive output, affect an individual's probability of survival, and are crucial in adapting an individual to its environmental surroundings. The sea anemone Metridium senile (L.) is highly variable in ecological distribution and life history, including rate of growth, individual size, and rate of asexual reproduction. Gonad size (measured as cross-sectional area of gonadal tissue) increases with body weight, so individuals should grow as large and as rapidly as possible to maximize individual sexual reproductive output. Cessation of growth and small body size in intertidal populations suggest that growth is constrained by genetic or environmental conditions. The growth of intertidal individuals transplanted to harbor-float panels demonstrated that growth limits are imposed by environmental factors, most probably limited food and feeding time and damage from wave exposure (which stimulates fragmentation). Individuals in harbor-float populations, which are continuously immersed, grow much larger, and large individuals comprise a greater proportion of the population than in the intertidal zone. The highest rate of fragmentation observed was on harbor-float panels. Patterns of growth and asexual reproduction provide adaptive advantages for M. senile. For harborfloat individuals, large individual size increases gamete production and may increase feeding efficiency. For intertidal individuals, asexual reproduction allows growth despite individual size constraints and rapid population growth, with specific advantages resulting from clone formation.     

Trade-offs between clonal and sexual reproduction in Sagittaria latifolia (Alismataceae) scale up to affect the fitness of entire clones

? Many plants combine sexual reproduction with vegetative propagation, but how trade-offs between these reproductive modes affect fitness is poorly understood. Although such trade-offs have been demonstrated at the level of individual shoots (ramets), there is little evidence that they scale up to affect genet fitness. For hermaphrodites, reproductive investment is further divided between female and male sexual functions. Female function should generally incur greater carbon costs than male function, which might involve greater nitrogen (N) costs. ? Using a common garden experiment with diclinous, clonal Sagittaria latifolia we manipulated investment in reproduction through female and male sex functions of 412 plants from monoecious and dioecious populations. ? We detected a 1?:?1 trade-off between biomass investment in female function and clonal reproduction. For male function, there was no apparent trade-off between clonal and sexual reproduction in terms of biomass investment. Instead, male function incurred a substantially higher N cost. ? Our results indicate that: trade-offs between investment in clonal propagation and sexual reproduction occur at the genet level in S.?latifolia; and sexual reproduction interferes with clonal expansion, with investment in female function limiting the quantity of clonal propagules produced, and investment in male function limiting the nutrient content of clonal propagules.     

Pubertal maturation is independent of family structure but daughters of divorced (but not dead) fathers start reproduction earlier

Several evolutionary hypotheses predict that girls growing up without a father present in the family mature and start reproduction earlier because father absence is a cue for environmental harshness and/or uncertainty that favours switching to a precocious life-history strategy. Most studies supporting these hypotheses have been performed in contemporary Western societies where the father absence is usually caused by divorce or abandonment. We performed a large retrospective cohort study in the mid-twentieth century Estonia, where different types of orphans and daughters of divorced fathers were exactly matched with girls from bi-parental families based on age, birth year, urban/rural origin and socioeconomic position of the family. Pubertal maturation, assessed on the basis of the breast development rate, did not associate with family structure. Daughters of divorced fathers started reproduction on average 9.2 months earlier than girls from bi-parental families and also tended to start reproduction earlier than girls whose fathers were dead. This finding is consistent with the view that fathers prone to divorce (and/or their spouses) differ from the rest of the population and also from the fathers prone to early death in some important characteristics that affect the reproductive rates of their daughters.     

Population genetics of complex life-cycle parasites: an illustration with trematodes

Accurate inferences on population genetics data require a sound underlying theoretical null model. Organisms alternating sexual and asexual reproduction during their life-cycle have been largely neglected in theoretical population genetic models, thus limiting the biological interpretation of population genetics parameters measured in natural populations. In this article, we derive the expectations of those parameters for the life-cycle of monoecious trematodes, a group comprising several important human and livestock parasites that obligatorily alternate sexual and asexual reproduction during their life-cycle. We model how migration rates between hosts, sexual and asexual mutation rates, adult selfing rate and the variance in reproductive success of parasites during the clonal phase affect the amount of neutral genetic diversity of the parasite (effective population size) and its apportionment within and between definitive hosts (using F-statistics). We demonstrate, in particular, that variance in reproductive success of clones, a parameter that has been completely overlooked in previous population genetics models, is very important in shaping the distribution of the genetic variability both within and among definitive hosts. Within definitive hosts, the parameter F(IS) (a measure of the deviation from random mating) is decreased by high variance in clonal reproductive success of larvae but increased by high adult self-fertilisation rates. Both clonal multiplication and selfing have similar effects on between-host genetic differentiation (F(ST)). Migration occurring before and after asexual reproduction can have different effects on the patterns of F(IS), depending on values of the other parameters such as the mutation rate. While the model applies to any hermaphroditic organism alternating sexual and clonal reproduction (e.g. many plants), the results are specifically discussed in the light of the limited population genetic data on monoecious trematodes available to date and their previous interpretation. We hope that our model will encourage more empirical population genetics studies on monoecious trematodes and other organisms with similar life-cycles.     

Effects of parasitic castration on plant resource allocation

It has been proposed that host castration is a parasite strategy to reallocate host resources from reproductive to vegetative functions to increase parasite fitness. Since resource partitioning between reproduction and vegetative growth can affect host life-history traits, parasite effects on resource allocation can affect both plant fitness and host-parasite coevolution. Field and greenhouse experiments were used to investigate the effects of host castration by the fungus Atkinsonella hypoxylon on the resource allocation and architecture of the grass Danthonia spicata. The results indicate that non-infected D. spicata can reallocate resources from reproduction to vegetative growth when resource allocation to reproduction is prevented. However, I found no evidence that fungal castration causes reallocation of resources from host reproduction to vegetative growth. Instead, infection reduces host biomass and the fungus directly utilizes resources that would have been used for host reproduction for its own reproduction. Received: 25 March 1999 / Accepted: 24 October 1999     

蚕豆蚜种群动态与蚕豆生理变化的关系

本文以蚕豆和蚕豆蚜构成的人工种间关系系统为对象,研究了蚜虫种群动态和植物生理变化的关系。发现蚕豆生理应激过程能影响蚕豆蚜种群的生殖率、存活率等种群特征,从而调节其种群的动态。蚕豆还能传导放大昆虫种群自主调节的信息。蚜虫种群的适应过程,包括减小种群数量(低生殖率和迁移),降低对植物的胁迫,从而维持种间关系系统的持续发展。本文还初步将植物的生理应激过程与昆虫种群的动态过程相耦联,建立动态模型,对种间关系的发展趋势进行了分析和讨论。     

孑遗植物四合木(Tetraena mongolica)的濒危肇因与机制

对我国特有单属种孑遗植物四合木 (Tetraena mongolica Maxim) 的地理分布、生境条件、种群数量动态、空间分布格局、种间关系、种群的生命表、生殖力表、有性生殖、无性繁殖和遗传变异等种群生态学特征及其濒危肇因进行了系统分析.研究结果表明:四合木种群为高群集分布,种群曲线属Leak凸型.年龄结构不合理,其存活曲线接近于Deevey ê型,且将演化为小种群,群落内的生境异质性显著,种群处于不稳定阶段.四合木从开花期到结果期,生殖分配 (RA) 呈下降的趋势,生殖过程中胚胎发生败育、种子向幼苗难以转化使其有性生殖受阻,生活史趋于断裂,是最终导致其濒危的重要内因.分布区城市化、工业化以及过度放牧等原因造成其种群孤立和生境破碎化是物种导致濒危的外因,四合木生态适应性和生境适宜性下降造成遗传多样性逐步丧失.同时提出对我国特有植物四合木进行异地保护的可能性与必然性.     

Effect of Adult Feeding on Male Mating Behaviour in the Butterfly, <Emphasis Type="Italic">Bicyclus anynana</Emphasis> (Lepidoptera: Nymphalidae)

Many species of lepidopterans supplement their nectar diet with foods rich in nitrogen and minerals, which are present only in trace amounts in nectar. We examined the effect of adult diet on mating behaviour and spermatophore characteristics in male Bicyclus anynana (Butler, 1879) butterflies, which feed on rotten fruits as adults. We found little effect of adult diet on male reproduction in terms of mating rate and sperm production, although males fed on fruit produced larger spermatophores on their first mating compared to males fed sugar only. We also examined how males allocate sperm across matings. Males ejaculate larger spermatophores during their first mating, and produce spermatophores containing decreasingly fewer non-fertile sperm with number of matings performed. Males that produced more non-fertile sperm on their first mating had reduced lifespan possibly indicating a trade-off between sperm production and adult longevity. It is suggested that adult diet has little affect on male ejaculate production and males feed on fruit to supplement their energetic carbon requirements.     

The bottleneck of JNK signaling: molecular and functional characteristics of MKK4 and MKK7

The functions of mitogen-activated protein kinases (MKKs) 4 and 7 are typically associated with the c-Jun N-terminal kinase (JNK) signaling pathway. Both MKKs synergistically phosphorylate different JNK isoforms and are therefore involved in numerous physiological (e.g. differentiation and proliferation) and pathological (e.g. apoptosis and tumorigenesis) processes. MKK4 and MKK7 share similar molecular characteristics as well as several upstream activators and scaffold proteins. However, their functions are non-redundant and determined by different stimuli, biochemical interactions and differential tissue distribution. The central question is how two MKKs regulate or affect the multiple actions of their JNK substrates. Similar to JNKs, MKK4 and MKK7 can simultaneously exert divergent functions in different cellular compartments and signalosomes. It is also important to realize that the MKK effects are splice variant-specific. The present review not only summarizes the various modes of MKK4 and MKK7 activation and activity, but also their functions. We also extensively describe their impact on JNK signaling, their molecular interactions resulting in the formation of context-specific signalosomes and the functional consequences of JNK deficiency.     

孑遗植物四合木(Tetraena mongolica)的濒危肇因与机制

对我国特有单属种孑遗植物四合木 (Tetraena mongolica Maxim) 的地理分布、生境条件、种群数量动态、空间分布格局、种间关系、种群的生命表、生殖力表、有性生殖、无性繁殖和遗传变异等种群生态学特征及其濒危肇因进行了系统分析。研究结果表明：四合木种群为高群集分布,种群曲线属Leak凸型。年龄结构不合理,其存活曲线接近于Deevey ê型,且将演化为小种群,群落内的生境异质性显著,种群处于不稳定阶段。四合木从开花期到结果期,生殖分配 (RA) 呈下降的趋势,生殖过程中胚胎发生败育、种子向幼苗难以转化使其有性生殖受阻,生活史趋于断裂,是最终导致其濒危的重要内因。分布区城市化、工业化以及过度放牧等原因造成其种群孤立和生境破碎化是物种导致濒危的外因,四合木生态适应性和生境适宜性下降造成遗传多样性逐步丧失。同时提出对我国特有植物四合木进行异地保护的可能性与必然性。     

Characterizing superspreading potential of infectious disease: Decomposition of individual transmissibility

In the context of infectious disease transmission, high heterogeneity in individual infectiousness indicates that a few index cases can generate large numbers of secondary cases, a phenomenon commonly known as superspreading. The potential of disease superspreading can be characterized by describing the distribution of secondary cases (of each seed case) as a negative binomial (NB) distribution with the dispersion parameter, k. Based on the feature of NB distribution, there must be a proportion of individuals with individual reproduction number of almost 0, which appears restricted and unrealistic. To overcome this limitation, we generalized the compound structure of a Poisson rate and included an additional parameter, and divided the reproduction number into independent and additive fixed and variable components. Then, the secondary cases followed a Delaporte distribution. We demonstrated that the Delaporte distribution was important for understanding the characteristics of disease transmission, which generated new insights distinct from the NB model. By using real-world dataset, the Delaporte distribution provides improvements in describing the distributions of COVID-19 and SARS cases compared to the NB distribution. The model selection yielded increasing statistical power with larger sample sizes as well as conservative type I error in detecting the improvement in fitting with the likelihood ratio (LR) test. Numerical simulation revealed that the control strategy-making process may benefit from monitoring the transmission characteristics under the Delaporte framework. Our findings highlighted that for the COVID-19 pandemic, population-wide interventions may control disease transmission on a general scale before recommending the high-risk-specific control strategies.     

Influence of the hyperparasitoid Dendrocerus carpenteri on the reproduction of the grain aphid Sitobion avenae

1.Alate S. avenae (F.) reproduced earlier and produced more offspring in the vicinity of Dendrocerus carpenteri (Curtis) females than control aphids without neighbouring hyperparasitoids.
2. This effect was evident only during the first 6 days of the aphid's reproduction period and did not affect the physiological condition of the aphids or their offspring.
3. Virgin hyperparasitoid females had a stronger influence on the reproduction rate of the aphids than did mated females, probably due to an unidentified emitted volatile sex pheromone.