A TEST OF THE CONJECTURE THAT G‐MATRICES ARE MORE STABLE THAN B‐MATRICES

The G‐matrix occupies an important position in evolutionary biology both as a summary of the inheritance of quantitative traits and as an ingredient in predicting how those traits will respond to selection and drift. Consequently, the stability of G has an important bearing on the accuracy of predicted evolutionary trajectories. Furthermore, G should evolve in response to stable features of the adaptive landscape and their trajectories through time. Although the stability and evolution of G might be predicted from knowledge of selection in natural populations, most empirical comparisons of G‐matrices have been made in the absence of such a priori predictions. We present a theoretical argument that within‐sex G‐matrices should be more stable than between‐sex B‐matrices because they are more powerfully exposed to multivariate stabilizing selection. We tested this conjecture by comparing estimates of B‐ and within‐sex G‐matrices among three populations of the garter snake Thamnophis elegans. Matrix comparisons using Flury's hierarchical approach revealed that within‐sex G‐matrices had four principal components in common (full CPC), whereas B‐matrices had only a single principal component in common and eigenvalues that were more variable among populations. These results suggest that within‐sex G is more stable than B , as predicted by our theoretical argument.     

Social evolution and the two elements of causation

The kin selection theory has recently been criticised on the basis of claiming that genetic relatedness does not play a causal role in the social evolution among individuals of insect societies. We outline here a line of criticism of this view by demonstrating two things. First, there are strong conceptual, theoretical and empirical reasons to think that close genetic relatedness has been necessary for the rise of the helper castes of social insects. And second, once we understand how causal explanation itself results from an interplay of two logically distinct elements, necessity and sufficiency, we can also understand the scenarios in which relatedness does not seem to play a causal role for evolution of helper castes. The result of this analysis is that we should be more careful about the way we frame the empirical data on the evolution of social behaviour.     

Mountain metacommunities: climate and spatial connectivity shape ant diversity in a complex landscape

Understanding what drives biodiversity patterns across scales is a central goal of ecology. Both environmental gradients and spatial landscape structure have been found to be important factors influencing species distributions and community composition, and partly reflect the balance of underlying deterministic and stochastic community processes. In some systems, environmental gradients and spatial connectivity are intertwined in that steep environmental gradients serve as boundaries on species movements and impose environment‐derived complex spatial structure to metacommunities. Mountainous landscapes are prime examples of this, and recent theory has linked principles of geomorphology, environmental gradients, and spatial structure to make predictions for resulting community patterns. In this context, we examine variation in taxonomic and phylogenetic ant diversity patterns along a geographic transect spanning > 5000 m in elevational range in the Hengduan mountains of southern China. We found that environmental gradients dominate variation in both alpha and beta diversity in this landscape, with alpha diversity strongly declining with elevation and beta diversity driven by elevational differences. However, within an elevational band spatial connectivity predicts beta diversity better than geographic distance. Our findings deviate from theoretical predictions in several ways, notably alpha diversity is monotonically declining and within‐band beta diversity is invariant with increasing elevation. The discrepancies between theory and observation may be explained by differences in the Hengduan landscape from idealized fluvial landscapes, such as a lack of a mid‐elevation peak in connectivity, as well as evolutionary limits on the source pool of species available to populate metacommunities at different elevations. The latter is supported by variation in phylogenetic community structure with elevation. Our results demonstrate the power of conceptual, statistical, and theoretical frameworks that integrate the roles of environment and spatial structure in metacommunities, but that additional work is needed to bridge the gap between abstract theory and real systems.     

Darwin's "beloved barnacles": tough lessons in variation

In 1846, burdened by insecurity and self-doubt, and having been convinced that he needed to study some group of organisms closely, Darwin embarked on an eight-year odyssey in the protean and perplexing world of barnacles. At the time, he was searching for evidence in support of his theory of evolution by natural selection. In the course of his long study of barnacles, however, he was not just validating his preexisting theoretical system, but was also modifying his views on such fundamental aspects as the universality of individual variation, which is the focus of this paper. According to this notion, the members of any population of living things are expected to exhibit sufficient differences from one another for natural selection to operate. By emphasizing the theoretical value of the barnacle project, my analysis contributes to the historiographic tradition which highlights the significance of the period between the first comprehensive formulation of the theory of evolution by natural selection in 1844 and its urgent publication in the late 1850s. In the course of these years, Darwin's theory was not just accumulating empirical laurels, but was also expected to adapt to a changing conceptual landscape.     

Global view of bionetwork dynamics: adaptive landscape

Based on recent work, I will give a nontechnical brief review of a powerful quantitative concept in biology, adaptive landscape, ini- tially proposed by S. Wright over 70 years ago, reintroduced by one of the founders of molecular biology and by others in different bio- logical contexts, but apparently forgotten by modem biologists for many years. Nevertheless, this concept finds an increasingly important role in the development of systems biology and bionetwork dynamics modeling, from phage lambda genetic switch to endogenous net- work for cancer genesis and progression. It is an ideal quantification to describe the robustness and stability of bionetworks. Here, I will first introduce five landmark proposals in biology on this concept, to demonstrate an important common thread in theoretical biology. Then I will discuss a few recent results, focusing on the studies showing theoretical consistency of adaptive landscape. From the perspec- tive of a working scientist and of what is needed logically for a dynamical theory when confronting empirical data, the adaptive landscape is useful both metaphorically and quantitatively, and has captured an essential aspect of biological dynamical processes. Though at the theoretical level the adaptive landscape must exist and it can be used across hierarchical boundaries in biology, many associated issues are indeed vague in their initial formulations and their quantitative realizations are not easy, and are good research topics for quantitative biologists. I will discuss three types of open problems associated with the adaptive landscape in a broader perspective.     

Evolutionary models of extended phenotypes

A variety of theoretical models incorporate phenotypes expressed in the external environment, but a core question is whether such traits generate dynamics that alter evolution. This has proven to be a challenging and controversial proposition. However, several recent modelling frameworks provide insight: indirect genetic effect (IGE) models, niche construction models, and evolutionary feedback models. These distinct approaches converge upon the observation that gene action at a distance generates feedback that expands the range of trait values and evolutionary rates that we should expect to observe in empirical studies. Such conceptual replication provides solid evidence that traits with extended effects have important evolutionary consequences, but more empirical work is needed to evaluate the predictive power of different modelling approaches.     

THE EVOLUTION OF LANDSCAPE GENETICS

The main objective of this special section is not to review the broad field of landscape genetics, but to provide a glimpse of how the developing landscape genetics perspective has the potential to change the way we study evolution. Evolutionary landscape genetics is the study of how migration and population structure affects evolutionary processes. As a field it dates back to Sewall Wright and the origin of theoretical population genetics, but empirical tests of adaptive processes of evolution in natural landscapes have been rare. Now, with recent developments in technology, methodology, and modeling tools, we are poised to trace adaptive genetic variation across space and through time. Not only will we see more empirical tests of classical theory, we can expect to see new phenomena emerging, as we reveal complex interactions among evolutionary processes as they unfold in natural landscapes.     

Quantitative genetic models of sexual selection

Modeling of R.A. Fisher's ideas about the evolution of male ornamentation using quantitative genetics began in the 1980s. Following an initial period of enthusiasm, interest in these models began to wane when theoretical studies seemed to show that the rapid evolution of ornaments would not occur if there were costs associated with female mate choice. Recent theoretical work has shown, however, that runaway evolution and other kinds of extensive diversification of ornaments and preferences can occur, even when female choice is costly. These new models highlight crucial parameters that profoundly influence evolutionary trajectories, but these parameters have been neglected in empirical studies. Here, we review quantitative genetic models of sexual selection with the aim of fostering communication and synergism between theoretical and empirical enterprises. We also point out several areas in which additional empirical work could distinguish between alternative models of evolution.     

The evolution of sex: empirical insights into the roles of epistasis and drift

Despite many years of theoretical and experimental work, the explanation for why sex is so common as a reproductive strategy continues to resist understanding. Recent empirical work has addressed key questions in this field, especially regarding rates of mutation accumulation in sexual and asexual organisms, and the roles of negative epistasis and drift as sources of adaptive constraint in asexually reproducing organisms. At the same time, new ideas about the evolution of sexual recombination are being tested, including intriguing suggestions of an important interplay between sex and genetic architecture, which indicate that sex and recombination could have affected their own evolution.     

The ecology of visual landscapes: Exploring the conceptual common ground of visual and ecological landscape indicators

This paper explores the conceptual common ground between visual and ecological landscape indicators. Indicators can be used to summarise complex information about landscape functions and the ability to extract a common set of indicators for analysing different landscape functions may provide valuable support for multiple-use planning.The development of landscape ecological indicators has been a very active research field that has resulted in a wide range of landscape metrics and composite indices with a strong conceptual base in landscape ecological principles. For the visual aspects of landscapes, however, this conceptual base is often missing and thus hindering progress in the development of indicators. This paper presents the results of an analysis of the correspondence between ecological indicators and visual indicators in order to explore whether there is common ground in both concept and operation. The conceptual level is presented for both ecological and visual indicators, as it is the concepts that define their aim and scope. We feel that transparency over this issue is crucial to the development and use of environmental indicators. The approach identified a candidate set of indicators that capture important aspects of both ecological and visual quality. The strength of the approach is that it forces us to focus on the identification of what we wish to indicate by means of landscape theory and assessment that are relevant to a specific landscape context. We believe that the approach presented here forms the basis for development of new methods for understanding the impact of landscape change in a management and planning context.     

Rethinking the conceptual foundations of habitat fragmentation research

The conceptual foundations of habitat fragmentation research have not kept pace with empirical advances in our understanding of species responses to landscape change, nor with theoretical advances in the wider disciplines of ecology. There is now real debate whether explicit recognition of ‘habitat fragmentation’ as an over‐arching conceptual domain will stimulate or hinder further progress toward understanding and mitigating the effects of landscape change. In this paper, we critically challenge the conceptual foundations of the discipline, and attempt to derive an integrated perspective on the best way to advance mechanistic understanding of fragmentation processes. We depict the inherent assumptions underlying the discipline as a ‘conceptual phase space’ of contrasting false dichotomies in fragmentation ‘problem space’. In our opinion, the key determinant of whether ‘habitat fragmentation’ can remain a cohesive framework lies in the concept of ‘interdependence’: 1) interdependence of landscape effects on species and 2) interdependence of species responses to landscape change. If there is non‐trivial interdependence among the various sub‐components of habitat fragmentation, or non‐trivial interdependence among species responses to landscape change, then there will be real heuristic value in ‘habitat fragmentation’ as a single conceptual domain. At present, the current paradigms entrenched in the fragmentation literature are implicitly founded on strict independence of landscape effects (e.g. the debate about the independent effects of habitat loss versus fragmentation per se) and strict independence of species responses (e.g. the individualistic species response models underpinning landscape continuum models), despite compelling evidence for interdependence in both effects and responses to fragmentation. We discuss how strong ‘interdependence’ of effects and responses challenges us to rethink long‐held views, and re‐cast the conceptual foundations of habitat fragmentation in terms of spatial context‐dependence in the effects of multiple interacting spatial components of fragmentation, and community context‐dependence in the responses of multiple interacting species to landscape change.     

Donating Embryos to Stem Cell Research The “Problem” of Gratitude

This paper is based on linked qualitative studies of the donation of human embryos to stem cell research carried out in the United Kingdom, Switzerland, and China. All three studies used semi-structured interview protocols to allow an in-depth examination of donors’ and non-donors’ rationales for their donation decisions, with the aim of gaining information on contextual and other factors that play a role in donor decisions and identifying how these relate to factors that are more usually included in evaluations made by theoretical ethics. Our findings have implications for one factor that has previously been suggested as being of ethical concern: the role of gratitude. Our empirical work shows no evidence that interpersonal gratitude is an important factor, but it does support the existence of a solidarity-based desire to “give something back” to medical research. Thus, we use empirical data to expand and refine the conceptual basis of bioethically theorizing the IVF–stem cell interface.     

13 RNA fitness landscapes

The origin of life is believed to have progressed through an RNA World, in which RNA acted as both genetic material and functional molecules. Understanding early evolution requires systematic knowledge of the relationship between RNA sequence and functional activity. In particular, knowing the structure of the fitness landscape of RNA is critical in estimating the probability of the emergence of functional sequences and the role of historical accident during evolution. Much theoretical work has been devoted to fitness landscapes, but experimental maps have been relatively limited. We use in vitro selection on a pool of short RNA sequences that nearly saturates sequence space to reconstruct the form of a comprehensive fitness landscape. We also study mutations during non-enzymatic polymerization to understand how early RNA replicators would ‘move’ in sequence space.     

Heterogeneous selection promotes maintenance of polymorphism in host–parasite interactions

Polymorphism in loci affecting host resistance and parasite virulence is characteristic for nearly all species and this genetic variation is considered to have profound consequences for the patterns of disease incidence, prevalence and evolution. The gene-for-gene (GFG) system is a well-characterized genetic interaction of host recognition and parasite antigenic loci for a wide range of plant-parasite interactions. Long-term maintenance of polymorphism in GFG systems has remained puzzling for both theoreticians and empiricists. Traditionally this diversity has been explained by tradeoffs with other life-history traits closely linked with fitness, yet empirical evidence for such costs has remained mixed. Here we argue that incorporating simple ecological reality – spatial structuring and gradient of environmental conditions – into host–parasite research will help us understand how polymorphism is maintained. While environmental conditions (biotic and abiotic factors) have been studied in depth in plant pathology for their influence on disease severity and plant yield, they have been rarely set into an evolutionary framework. We briefly review recent data on natural plant–parasite metapopulations and theoretical models moving from single population models towards metapopulation theory to reveal in just how many ways spatial structuring may affect the coevolutionary process. We clarify also how spatially heterogeneous selection, through G×E (or G×G×E) interactions, may be particularly important for natural host–parasite interactions and suggest that this provides the unifying ground upon which future theoretical and empirical work should be build on.     

Beyond the fragmentation threshold hypothesis: regime shifts in biodiversity across fragmented landscapes

Ecological systems are vulnerable to irreversible change when key system properties are pushed over thresholds, resulting in the loss of resilience and the precipitation of a regime shift. Perhaps the most important of such properties in human-modified landscapes is the total amount of remnant native vegetation. In a seminal study Andrén proposed the existence of a fragmentation threshold in the total amount of remnant vegetation, below which landscape-scale connectivity is eroded and local species richness and abundance become dependent on patch size. Despite the fact that species patch-area effects have been a mainstay of conservation science there has yet to be a robust empirical evaluation of this hypothesis. Here we present and test a new conceptual model describing the mechanisms and consequences of biodiversity change in fragmented landscapes, identifying the fragmentation threshold as a first step in a positive feedback mechanism that has the capacity to impair ecological resilience, and drive a regime shift in biodiversity. The model considers that local extinction risk is defined by patch size, and immigration rates by landscape vegetation cover, and that the recovery from local species losses depends upon the landscape species pool. Using a unique dataset on the distribution of non-volant small mammals across replicate landscapes in the Atlantic forest of Brazil, we found strong evidence for our model predictions--that patch-area effects are evident only at intermediate levels of total forest cover, where landscape diversity is still high and opportunities for enhancing biodiversity through local management are greatest. Furthermore, high levels of forest loss can push native biota through an extinction filter, and result in the abrupt, landscape-wide loss of forest-specialist taxa, ecological resilience and management effectiveness. The proposed model links hitherto distinct theoretical approaches within a single framework, providing a powerful tool for analysing the potential effectiveness of management interventions.     

Complexity and evolution: What everybody knows

The consensus among evolutionists seems to be (and has been for at least a century) that the morphological complexity of organisms increases in evolution, although almost no empirical evidence for such a trend exists. Most studies of complexity have been theoretical, and the few empirical studies have not, with the exception of certain recent ones, been especially rigorous; reviews are presented of both the theoretical and empirical literature. The paucity of evidence raises the question of what sustains the consensus, and a number of suggestions are offered, including the possibility that certain cultural and/or perceptual biases are at work. In addition, a shift in emphasis from theoretical to empirical inquiry is recommended for the study of complexity, and guidelines for future empirical studies are proposed.     

On the role of spatial stochastic models in understanding landscape indices in ecology

Spatial stochastic models play an important role in understanding and predicting the behaviour of complex systems. Such models may be implemented with explicit knowledge of only a limited number of parameters relating to spatial relationships among locations. Consequently, they are often used instead of deterministic‐mechanistic models, which may potentially require an unrealistically large number of parameters. Currently, in contrast to spatial stochastic models, the parameterization of the joint spatial distribution of objects in landscape models is more often implicit than explicit. Here, we investigate the similarities and differences between bona fide spatial stochastic models and landscape models by focusing mostly on the relationships between processes, their realizations (patterns), representation and measurement, and their use in exploratory as well as confirmatory data analysis. One of the most important outcomes of recognizing the importance of stochastic processes is the acknowledgement that the spatial pattern observed in a landscape is only one realization of that process. Hence, while ecologists have been using landscape pattern indices (LPIs) to characterize landscape heterogeneity and/or make inferences about processes shaping the landscape, no stochastic modelling framework has been developed for their proper statistical elucidation. Consequently, several (mis)uses of LPIs draw conclusions about landscapes which are suspect. We show that several reports about sensitivities of LPIs to measurements have common roots that can be made explicitly manageable by adopting stochastic models of spatial structure. The key parameters of these stochastic models are composition and configuration, which, in general, cannot be estimated independently from each other. We outline how to develop the stochastic framework to interpret observations and make some recommendations to practitioners about everyday usage. The conceptual linkages between patterns and processes are particularly important in light of recent efforts to bridge the static‐structural and the dynamic‐analytic traditions of ecology.     

Genetic and environment-induced pathways to innovation: on the possibility of a universal relationship between robustness and adaptation in complex biological systems

Recently, there has been considerable interest in the idea that mutational robustness enhances the propensity for future adaptations, i.e. evolvability, if evolution proceeds over a neutral network that extends far throughout a fitness landscape. While the genetic neutral network (NN-G) model may have important implications to our understanding of evolution, little has been done to integrate these theoretical developments with empirical evidence that heritable phenotypes can also originate and become fixated as a result of changes in the environment. In this brief commentary, I reconsider the role of environmental change in the adaptation of species and ask whether positive robustness-evolvability relationships might exist not only for genetic but also environmental buffering. In particular, I ask whether the insensitivity of species fitness towards variability in its environment can have a positive influence on the likelihood of future environment-induced adaptations (i.e. ecological opportunities) in a manner analogous to that proposed by the NN-G model. After outlining scenarios where such a counter-intuitive relationship appears plausible, I comment on the merits of evolutionary theories that can integrate complementary pathways to adaptation under static and time-variant environments. I also speculate on some of the features that such a theory might have.     

Is there such a thing as landscape genetics?

For a scientific discipline to be interdisciplinary, it must satisfy two conditions; it must consist of contributions from at least two existing disciplines, and it must be able to provide insights, through this interaction, that neither progenitor discipline could address. In this study, I examine the complete body of peer‐reviewed literature self‐identified as landscape genetics (LG) using the statistical approaches of text mining and natural language processing. The goal here was to quantify the kinds of questions being addressed in LG studies, the ways in which questions are evaluated mechanistically, and how they are differentiated from the progenitor disciplines of landscape ecology and population genetics. I then circumscribe the main factions within published LG studies examining the extent to which emergent questions are being addressed and highlighting a deep bifurcation between existing individual‐ and population‐based approaches. I close by providing some suggestions on where theoretical and analytical work is needed if LGs is to serve as a real bridge connecting evolution and ecology sensu lato.