Test for latitudinal variation of life history,behavior and mortality in the strictly univoltine damselfly Sympecma fusca (Zygoptera: Lestidae)

A wealth of evidence shows that combinations of ecological stressors interact in shaping life history traits, but little is known about how ecological stressors combine with different seasonal time constraints to shape life history, behavior and mortality across populations. We studied life history, behavior and mortality rate in two latitudinally distant populations of the strictly univoltine, adult‐overwintering damselfly Sympecma fusca. Results from laboratory common‐garden and outdoor experiments indicated countergradient variation of larval development time and growth rate: the more time‐constrained larvae showed faster development and a higher growth rate. This finding led to larger size at emergence in the more time‐constrained individuals. Under conditions of intraspecific interaction (outdoor experiment), northern individuals showed lower survival than southern ones, presumably due to cannibalism. In the absence of intraspecific interactions (laboratory experiment), northern and southern larvae did not differ in survival. Finally, laboratory‐grown northern and southern larvae did not differ in activity level. This is the first time that compensation for seasonal time constraints has been shown in a temperate odonate species that overwinters in the adult stage.     

Ecological constraints, life history traits and the evolution of cooperative breeding

The ecological constraints hypothesis is widely accepted as an explanation for the evolution of delayed dispersal in cooperatively breeding birds. Intraspecific studies offer the strongest support. Observational studies have demonstrated a positive association between the severity of ecological constraints and the prevalence of cooperation, and experimental studies in which constraints on independent breeding were relaxed resulted in helpers moving to adopt the vacant breeding opportunities. However, this hypothesis has proved less successful in explaining why cooperative breeding has evolved in some species or lineages but not in others. Comparative studies have failed to identify ecological factors that differ consistently between cooperative and noncooperative species. The life history hypothesis, which emphasizes the role of life history traits in the evolution of cooperative breeding, offers a solution to this difficulty. A recent analysis showed that low adult mortality and low dispersal predisposed certain lineages to show cooperative behaviour, given the right ecological conditions. This represents an important advance, not least by offering an explanation for the patchy phylogenetic distribution of cooperative breeding. We discuss the complementary nature of these two hypotheses and suggest that rather than regarding life history traits as predisposing and ecological factors as facilitating cooperation, they are more likely to act in concert. While acknowledging that different cooperative systems may be a consequence of different selective pressures, we suggest that to identify the key differences between cooperative and noncooperative species, a broad constraints hypothesis that incorporates ecological and life history traits in a single measure of 'turnover of breeding opportunities' may provide the most promising avenue for future comparative studies. Copyright 2000 The Association for the Study of Animal Behaviour.     

Community assembly, stability and signatures of dynamical constraints on food web structure

To understand the dynamics of natural species communities, a major challenge is to quantify the relationship between their assembly, stability, and underlying food web structure. To this end, two complementary aspects of food web structure can be related to community stability: sign structure, which refers to the distributions of trophic links irrespective of interaction strengths, and interaction strength structure, which refers to the distributions of interaction strengths with or without consideration of sign structure. In this paper, using data from a set of relatively well documented community food webs, I show that natural communities generally exhibit a sign structure that renders their stability sensitive to interaction strengths. Using a Lotka-Volterra type population dynamical model, I then show that in such communities, individual consumer species with high values of a measure of their total biomass acquisition rate, which I term “weighted generality”, tend to undermine community stability. Thus consumer species’ trophic modules (a species and all its resource links) should be “selected” through repeated immigrations and extinctions during assembly into configurations that increase the probability of stable coexistence within the constraints of the community's trophic sign structure. The presence of such constraints can be detected by the incidence and strength of certain non-random structural characteristics. These structural signatures of dynamical constraints are readily measurable, and can be used to gauge the importance of interaction-driven dynamical constraints on communities during and after assembly in natural communities.     

Game Dynamics with Learning and Evolution of Universal Grammar

We investigate a model of language evolution, based on population game dynamics with learning. First, we examine the case of two genetic variants of universal grammar (UG), the heart of the human language faculty, assuming each admits two possible grammars. The dynamics are driven by a communication game. We prove using dynamical systems techniques that if the payoff matrix obeys certain constraints, then the two UGs are stable against invasion by each other, that is, they are evolutionarily stable. Then, we prove a similar theorem for an arbitrary number of disjoint UGs. In both theorems, the constraints are independent of the learning process. Intuitively, if a mutation in UG results in grammars that are incompatible with the established languages, then the mutation will die out because mutants will be unable to communicate and therefore unable to realize any potential benefit of the mutation. An example for which these theorems do not apply shows that compatible mutations may or may not be able to invade, depending on the population's history and the learning process. These results suggest that the genetic history of language is constrained by the need for compatibility and that mutations in the language faculty may have died out or taken over due more to historical accident than to any straightforward notion of relative fitness. MSC 1991: 37N25 · 92D15 · 91F20     

Life history plasticity to combined time and biotic constraints in Lestes damselflies from vernal and temporary ponds

Little is known about the ecological correlates and macro-evolution of life history plasticity to time constraints. Here, we compared age and mass at emergence and their plasticity toward combinations of time constraints (delayed larval development), food level and predation risk (caged dragonfly larvae) by rearing two temporary-pond Lestes damselflies ( L. congener and L. forcipatus ) and the derived vernal-pond L. dryas from the eggs until adult emergence in outdoor tubs. Life history plasticity under time constraints, low food and predation risk was as predicted by optimality models. Delayed larvae in all three species accelerated development and showed a lower fat content and a substantially elevated mortality rate. At low food, all species emerged later at a smaller mass. Also under predation risk adults of all species emerged later, and (at high food) at a smaller mass. Unexpectedly, delayed larvae did not show a smaller life history response to predation risk imposed by dragonfly larvae. Compared to the two temporary-pond Lestes studied, the derived species that invaded more ephemeral vernal ponds showed a faster development rate and a lower deceleration of development to low food. It also showed a lower acceleration of development to time constraints, possibly reflecting that it reached development rates near to its physiological maximum. Unexpectedly, the vernal-pond Lestes did not slow its development less under predation risk. Our results stress the importance of evaluating ecological and evolutionary correlates of life history plasticity under as realistic conditions as possible.     

Using multiple interdependency to separate functional from phylogenetic correlations in protein alignments

MOTIVATION: Multiple sequence alignments of homologous proteins are useful for inferring their phylogenetic history and to reveal functionally important regions in the proteins. Functional constraints may lead to co-variation of two or more amino acids in the sequence, such that a substitution at one site is accompanied by compensatory substitutions at another site. It is not sufficient to find the statistical correlations between sites in the alignment because these may be the result of several undetermined causes. In particular, phylogenetic clustering will lead to many strong correlations. RESULTS: A procedure is developed to detect statistical correlations stemming from functional interaction by removing the strong phylogenetic signal that leads to the correlations of each site with many others in the sequence. Our method relies upon the accuracy of the alignment but it does not require any assumptions about the phylogeny or the substitution process. The effectiveness of the method was verified using computer simulations and then applied to predict functional interactions between amino acids in the Pfam database of alignments.     

A guide to practical babooning: Historical,social, and cognitive contingency

As ecologically adaptable animals, baboons are distributed widely across Africa, and display a variety of morphological and behavioral differences that reflect both local ecology and a complex evolutionary history. As long‐lived, slowly reproducing animals, baboons face numerous ecological challenges to survival and successful reproduction. As group‐living animals, the social world presents an equally diverse array of challenges that require the negotiation of individual needs within the constraints imposed by others. Understanding how all these facets of baboon evolutionary history, life history, ecology, sociality, and cognition fit together is an enormous but engaging challenge, and despite one hundred years of study, it is clear there is a still much to learn about the various natural histories of baboons. What also is clear, however, is that an appreciation of contingency holds the key to understanding all these facets of baboon evolution and behavior. In what follows, I hope to illustrate exactly what I mean by this, highlighting along the way that history is not to be ignored, variability is information and not merely “noise”, and that behavioral and cognitive complexity can be two very different things.     

Understanding the role of floral development in the evolution of angiosperm flowers: clarifications from a historical and physico-dynamic perspective

Flower morphology results from the interaction of an established genetic program, the influence of external forces induced by pollination systems, and physical forces acting before, during and after initiation. Floral ontogeny, as the process of development from a meristem to a fully developed flower, can be approached either from a historical perspective, as a “recapitulation of the phylogeny” mainly explained as a process of genetic mutations through time, or from a physico-dynamic perspective, where time, spatial pressures, and growth processes are determining factors in creating the floral morphospace. The first (historical) perspective clarifies how flower morphology is the result of development over time, where evolutionary changes are only possible using building blocks that are available at a certain stage in the developmental history. Flowers are regulated by genetically determined constraints and development clarifies specific transitions between different floral morphs. These constraints are the result of inherent mutations or are induced by the interaction of flowers with pollinators. The second (physico-dynamic) perspective explains how changes in the physical environment of apical meristems create shifts in ontogeny and this is reflected in the morphospace of flowers. Changes in morphology are mainly induced by shifts in space, caused by the time of initiation (heterochrony), pressure of organs, and alterations of the size of the floral meristem, and these operate independently or in parallel with genetic factors. A number of examples demonstrate this interaction and its importance in the establishment of different floral forms. Both perspectives are complementary and should be considered in the understanding of factors regulating floral development. It is suggested that floral evolution is the result of alternating bursts of physical constraints and genetic stabilization processes following each other in succession. Future research needs to combine these different perspectives in understanding the evolution of floral systems and their diversification.     

Cannibalism-mediated life history plasticity to combined time and food stress

There is increasing awareness that combinations of biotic and time stress interact in shaping life history plasticity. Despite being widespread and abundant, the role of cannibalism in linking both types of constraints to life history plasticity has been largely neglected. Moreover, no studies disentangled direct (due to the extra meal) and indirect (due to the elimination of the competitor) life history effects of cannibalism, and little is known about their differential dependency on these constraints. We studied effects of cannibalism on the life history of the damselfly Lestes viridis under combinations of time stress (by manipulating the perceived time available in the growth season) and food stress. We reared larvae per two and disentangled direct and indirect effects of cannibalism by preventing cannibalism in half of the cups and by manipulating the per capita food increase after cannibalism. Cannibalism was more frequent under both time stress and food stress and our results show it may help cannibals to compensate for the negative effects of these constraints imposed on life history. Both direct and indirect benefits of cannibalism (increased development and growth rates, larger mass at emergence) were dependent on the timing of cannibalism, being more pronounced or only present when cannibalism occurred early. Moreover, we found that the ecological constraints (time stress and food stress) also differentially shaped some of the direct and indirect effects. Given the differential context-dependency of direct and indirect effects and the fact that direct and indirect life history effects may be both important in shaping life history, disentangling these effects is critical to mechanistically understand under which conditions cannibalism is expected to be adaptive or not.     

Quantitative genetics of migration syndromes: a study of two barn swallow populations

Migration is a complex trait although little is known about genetic correlations between traits involved in such migration syndromes. To assess the migratory responses to climate change, we need information on genetic constraints on evolutionary potential of arrival dates in migratory birds. Using two long-term data sets on barn swallows Hirundo rustica (from Spain and Denmark), we show for the first time in wild populations that spring arrival dates are phenotypically and genetically correlated with morphological and life history traits. In the Danish population, length of outermost tail feathers and wing length were negatively genetically correlated with arrival date. In the Spanish population, we found a negative genetic correlation between arrival date and time elapsed between arrival date and laying date, constraining response to selection that favours both early arrival and shorter delays. This results in a decreased rate of adaptation, not because of constraints on arrival date, but constraints on delay before breeding, that is, a trait that can be equally important in the context of climate change.     

Flexible life history responses to flower and rosette bud removal in three perennial herbs

In a garden experiment we investigated the response to continuous removal of either flower buds or rosette buds in three perennial grassland species ( Hypochaeris radicata , Succisa pratensis and Centaurea jacea ), which differ in longevity and flowering type. We distinguished two possible responses: compensation for lost buds by making more buds of the same type, and switching towards development of other life history functions. Both responses were demonstrated in our experiment, but bud removal had significantly different effects in each of the three species. The degree of compensation and the expression of trade-offs between life history functions differed markedly between species and seem related to longevity and developmental constraints. With respect to switching, our results suggest costs of reproduction and a trade-off between life history functions, at least for Hypochaeris and Succisa . For these species weight of new rosettes increased when resource allocation to flowering was inhibited. In Hypochaeris, we see that both compensation for lost flower buds and switching from lost rosette buds increased production of flower buds, underscoring the pivotal role of sexual reproduction in this short-lived species. The most prominent response seen in Centaurea is compensation for lost rosette buds, indicating that this long-lived species with monocarpic rosettes relies on rosette formation. Although Succisa does respond to bud removal, time is an important constraint in this species with long-lived rosettes and preformed flowering stalks. Trade-offs in Succisa seem to operate at a larger time scale, requiring long-lasting experiments to reveal them. We conclude that the response of these species to inflicted damage is likely to be linked to their longevity and developmental constraints.     

Testing multiple coordination constraints with a novel bimanual visuomotor task

The acquisition of a new bimanual skill depends on several motor coordination constraints. To date, coordination constraints have often been tested relatively independently of one another, particularly with respect to isofrequency and multifrequency rhythms. Here, we used a new paradigm to test the interaction of multiple coordination constraints. Coordination constraints that were tested included temporal complexity, directionality, muscle grouping, and hand dominance. Twenty-two healthy young adults performed a bimanual dial rotation task that required left and right hand coordination to track a moving target on a computer monitor. Two groups were compared, either with or without four days of practice with augmented visual feedback. Four directional patterns were tested such that both hands moved either rightward (clockwise), leftward (counterclockwise), inward or outward relative to each other. Seven frequency ratios (3∶1, 2∶1, 3∶2, 1∶1, 2∶3. 1∶2, 1∶3) between the left and right hand were introduced. As expected, isofrequency patterns (1∶1) were performed more successfully than multifrequency patterns (non 1∶1). In addition, performance was more accurate when participants were required to move faster with the dominant right hand (1∶3, 1∶2 and 2∶3) than with the non-dominant left hand (3∶1, 2∶1, 3∶2). Interestingly, performance deteriorated as the relative angular velocity between the two hands increased, regardless of whether the required frequency ratio was an integer or non-integer. This contrasted with previous finger tapping research where the integer ratios generally led to less error than the non-integer ratios. We suggest that this is due to the different movement topologies that are required of each paradigm. Overall, we found that this visuomotor task was useful for testing the interaction of multiple coordination constraints as well as the release from these constraints with practice in the presence of augmented visual feedback.     

On sex, mate selection and the red queen.

The widespread occurrence of sexual reproduction despite the two-fold disadvantage of producing males, is still an unsolved mystery in evolutionary biology. One explanatory theory, called the "Red Queen" hypothesis, states that sex is an adaptation to escape from parasites. A more recent hypothesis, the mate selection hypothesis, assumes that non-random mating, possible only with sex, accelerates the evolution of beneficial traits. This paper tests these two hypotheses, using an agent-based or "micro-analytic" evolutionary algorithm where host-parasite interaction is simulated adhering to biological reality. While previous simpler models testing the "Red Queen" hypothesis considered mainly haploid hosts, stable population density, random mating and simplified expression of fitness, our more realistic model allows diploidy, mate selection, live history constraints and variable population densities. Results suggest that the Red Queen hypothesis is not valid for more realistic evolutionary scenarios and that each of the two hypotheses tested seem to explain partially but not exhaustively the adaptive value of sex. Based on the results we suggest that sexual populations in nature should avoid both, maximizing outbreeding or maximizing inbreeding and should acquire mate selection strategies which favour optimal ranges of genetic mixing in accordance with environmental challenges.     

Viruses' Life History: Towards a Mechanistic Basis of a Trade-Off between Survival and Reproduction among Phages

Life history theory accounts for variations in many traits involved in the reproduction and survival of living organisms, by determining the constraints leading to trade-offs among these different traits. The main life history traits of phages—viruses that infect bacteria—are the multiplication rate in the host, the survivorship of virions in the external environment, and their mode of transmission. By comparing life history traits of 16 phages infecting the bacteria Escherichia coli, we show that their mortality rate is constant with time and negatively correlated to their multiplication rate in the bacterial host. Even though these viruses do not age, this result is in line with the trade-off between survival and reproduction previously observed in numerous aging organisms. Furthermore, a multiple regression shows that the combined effects of two physical parameters, namely, the capsid thickness and the density of the packaged genome, account for 82% of the variation in the mortality rate. The correlations between life history traits and physical characteristics of virions may provide a mechanistic explanation of this trade-off. The fact that this trade-off is present in this very simple biological situation suggests that it might be a fundamental property of evolving entities produced under constraints. Moreover, such a positive correlation between mortality and multiplication reveals an underexplored trade-off in host–parasite interactions.     

Evolution of dispersal and life history interact to drive accelerating spread of an invasive species

Populations on the edge of an expanding range are subject to unique evolutionary pressures acting on their life‐history and dispersal traits. Empirical evidence and theory suggest that traits there can evolve rapidly enough to interact with ecological dynamics, potentially giving rise to accelerating spread. Nevertheless, which of several evolutionary mechanisms drive this interaction between evolution and spread remains an open question. We propose an integrated theoretical framework for partitioning the contributions of different evolutionary mechanisms to accelerating spread, and we apply this model to invasive cane toads in northern Australia. In doing so, we identify a previously unrecognised evolutionary process that involves an interaction between life‐history and dispersal evolution during range shift. In roughly equal parts, life‐history evolution, dispersal evolution and their interaction led to a doubling of distance spread by cane toads in our model, highlighting the potential importance of multiple evolutionary processes in the dynamics of range expansion.     

Viruses' Life History: Towards a Mechanistic Basis of a Trade-Off between Survival and Reproduction among Phages

Life history theory accounts for variations in many traits involved in the reproduction and survival of living organisms, by determining the constraints leading to trade-offs among these different traits. The main life history traits of phages—viruses that infect bacteria—are the multiplication rate in the host, the survivorship of virions in the external environment, and their mode of transmission. By comparing life history traits of 16 phages infecting the bacteria Escherichia coli, we show that their mortality rate is constant with time and negatively correlated to their multiplication rate in the bacterial host. Even though these viruses do not age, this result is in line with the trade-off between survival and reproduction previously observed in numerous aging organisms. Furthermore, a multiple regression shows that the combined effects of two physical parameters, namely, the capsid thickness and the density of the packaged genome, account for 82% of the variation in the mortality rate. The correlations between life history traits and physical characteristics of virions may provide a mechanistic explanation of this trade-off. The fact that this trade-off is present in this very simple biological situation suggests that it might be a fundamental property of evolving entities produced under constraints. Moreover, such a positive correlation between mortality and multiplication reveals an underexplored trade-off in host–parasite interactions.     

Did surface temperatures constrain microbial evolution?

The proposition that glaciation may not have occurred before the Cenozoic--albeit not yet a consensus position--nevertheless raises for reconsideration the surface temperature history of the earth. Glacial episodes, from the Huronian (2.3 billion years ago; BYA) through the late Paleozoic (320 to 250 million years ago; MYA) have been critical constraints on estimation of the upper bounds of temperature (Crowley 1983, Kasting and Toon 1989). Once removed, few if any constraints on the upper temperature limit other than life remain. Walker (1982) recognized that life provides an upper limit to temperature in the Precambrian. We propose a more radical concept: the upper temperature limit for viable growth of a given microbial group corresponds to the actual surface temperature at the time of the group's first appearance. In particular, we propose here that two major evolutionary developments--the emergence of cyanobacteria and aerobic eukaryotes--can be used to determine surface temperature in the Precambrian, and that only subsequent cooling mediated by higher plants and then angiosperms permitted what may possibly be the earth's first glaciation in the late Cenozoic.     

Phenotypic Plasticity in the Annual Weed Polygonum aviculare

Reaction norms of fourteen life history and morphological traits were investigated in four tetra- and two hexaploid genotypes of the annual weed species complex, Polygonum aviculare. The plants were cultivated in six treatments consisting of factorial combinations of three pot sizes and two fertility levels. All characters, except life span, were plastic but the relative importance of genotype (G), treatment (T) and interaction (G × T) to total variance was strongly trait-specific. Consistent genetic differentiation, not correlated with ploidy level, was found in metamer size and life history: genotypes originating from trampled sites had smaller metamers and shorter shoots while those originating from sites with a short growing season, due to weeding activities, had a shorter life span, an earlier flowering date and a higher biomass allocation to reproduction compared to genotypes from less disturbed sites. Significant variation was found in reaction norms for all characters, including a lower amount of plasticity in metamer size in genotypes with numerous metamers and a lower amount of plasticity in total weight in shortlived genotypes. This suggested that variation in phenotypic plasticity reflected developmental constraints imposed by contrasting life span and metamer size in different genotypes. There was no evidence for niche differentiation along the soil resource gradient, suggesting that the species is comprised of “general purpose” genotypes with respect to soil fertility. It is concluded that the Polygonum aviculare complex has evolved a “dual” adaptive strategy i.e. a combination of genetic polymorphism and high phenotypic plasticity.     

Thermodynamic constraints on the assembly and diversity of microbial ecosystems are different near to and far from equilibrium

Non-equilibrium thermodynamics has long been an area of substantial interest to ecologists because most fundamental biological processes, such as protein synthesis and respiration, are inherently energy-consuming. However, most of this interest has focused on developing coarse ecosystem-level maximisation principles, providing little insight into underlying mechanisms that lead to such emergent constraints. Microbial communities are a natural system to decipher this mechanistic basis because their interactions in the form of substrate consumption, metabolite production, and cross-feeding can be described explicitly in thermodynamic terms. Previous work has considered how thermodynamic constraints impact competition between pairs of species, but restrained from analysing how this manifests in complex dynamical systems. To address this gap, we develop a thermodynamic microbial community model with fully reversible reaction kinetics, which allows direct consideration of free-energy dissipation. This also allows species to interact via products rather than just substrates, increasing the dynamical complexity, and allowing a more nuanced classification of interaction types to emerge. Using this model, we find that community diversity increases with substrate lability, because greater free-energy availability allows for faster generation of niches. Thus, more niches are generated in the time frame of community establishment, leading to higher final species diversity. We also find that allowing species to make use of near-to-equilibrium reactions increases diversity in a low free-energy regime. In such a regime, two new thermodynamic interaction types that we identify here reach comparable strengths to the conventional (competition and facilitation) types, emphasising the key role that thermodynamics plays in community dynamics. Our results suggest that accounting for realistic thermodynamic constraints is vital for understanding the dynamics of real-world microbial communities.