Reducing ambiguity in describing plant–insect interactions: "preference", "acceptability" and "electivity"

Ecologists and evolutionary biologists have a common interest in plant–insect interactions. Ecologists develop terminology describing patterns of association between plants and insects, while evolutionary biologists use the same words to denote potentially heritable traits of individuals. Use of the same terms to describe both traits of the interaction and traits of the organisms hinders communication. An example is "preference", often used by ecologists to denote properties of the plant–insect interaction and by behavioural or evolutionary biologists to denote insect traits. The existing term "electivity" could be incorporated into the lexicon of plant–insect interactions to supplant the ecological use of "preference". The term "preference" would then denote a behavioural trait of the insect. The mirror-image trait of the plant would be "acceptability". This could be a step towards a common terminology that would be usable by both ecologists and evolutionists.     

Developmental system drift and flexibility in evolutionary trajectories

SUMMARY The comparative analysis of homologous characters is a staple of evolutionary developmental biology and often involves extrapolating from experimental data in model organisms to infer developmental events in non-model organisms. In order to determine the general importance of data obtained in model organisms, it is critical to know how often and to what degree similar phenotypes expressed in different taxa are formed by divergent developmental processes. Both comparative studies of distantly related species and genetic analysis of closely related species indicate that many characters known to be homologous between taxa have diverged in their morphogenetic or gene regulatory underpinnings. This process, which we call "developmental system drift" (DSD), is apparently ubiquitous and has significant implications for the flexibility of developmental evolution of both conserved and evolving characters. Current data on the population genetics and molecular mechanisms of DSD illustrate how the details of developmental processes are constantly changing within evolutionary lineages, indicating that developmental systems may possess a great deal of plasticity in their responses to natural selection.     

Evolutionary theory of bacterial quorum sensing: when is a signal not a signal?

The term quorum sensing (QS) is used to describe the communication between bacterial cells, whereby a coordinated population response is controlled by diffusible molecules produced by individuals. QS has not only been described between cells of the same species (intraspecies), but also between species (interspecies) and between bacteria and higher organisms (inter-kingdom). The fact that QS-based communication appears to be widespread among microbes is strange, considering that explaining both cooperation and communication are two of the greatest problems in evolutionary biology. From an evolutionary perspective, intraspecies signalling can be explained using models such as kin selection, but when communication is described between species, it is more difficult to explain. It is probable that in many cases this involves QS molecules being used as 'cues' by other species as a guide to future action or as manipulating molecules whereby one species will 'coerce' a response from another. In these cases, the usage of QS molecules cannot be described as signalling. This review seeks to integrate the evolutionary literature on animal signalling with the microbiological literature on QS, and asks whether QS within bacteria is true signalling or whether these molecules are also used as cues or for the coercion of other cells.     

Pathogen pressure puts immune defense into perspective

The extent to which organisms can protect themselves from disease depends on both the immune defenses they maintain and the pathogens they face. At the same time, immune systems are shaped by the antigens they encounter, both over ecological and evolutionary time. Ecological immunologists often recognize these interactions, yet ecological immunology currently lacks major efforts to characterize the environmental, host-independent, antigenic pressures to which all animals are exposed. Failure to quantify relevant diseases and pathogens in studies of ecological immunology leads to contradictory hypotheses. In contrast, including measures of environmental and host-derived commensals, pathogens, and other immune-relevant organisms will strengthen the field of ecological immunology. In this article, we examine how pathogens and other organisms shape immune defenses and highlight why such information is essential for a better understanding of the causes of variation in immune defenses. We introduce the concept of "operative protection" for understanding the role of immunologically relevant organisms in shaping immune defense profiles, and demonstrate how the evolutionary implications of immune function are best understood in the context of the pressures that diseases and pathogens bring to bear on their hosts. We illustrate common mistakes in characterizing these immune-selective pressures, and provide suggestions for the use of molecular and other methods for measuring immune-relevant organisms.     

Tactics of evasion: strategies used by signallers to deter eavesdropping enemies from exploiting communication systems

Eavesdropping predators, parasites and parasitoids exploit signals emitted by their prey and hosts for detection, assessment, localization and attack, and in the process impose strong selective pressures on the communication systems of the organisms they exploit. Signallers have evolved numerous anti-eavesdropper strategies to mitigate the trade-off between the costs imposed from signal exploitation and the need for conspecific communication. Eavesdropper strategies fall along a continuum from opportunistic to highly specialized, and the tightness of the eavesdropper–signaller relationship results in differential pressures on communication systems. A wide variety of anti-eavesdropper strategies mitigate the trade-off between eavesdropper exploitation and conspecific communication. Antagonistic selection from eavesdroppers can result in diverse outcomes including modulation of signalling displays, signal structure, and evolutionary loss or gain of a signal from a population. These strategies often result in reduced signal conspicuousness and in decreased signal ornamentation. Eavesdropping enemies, however, can also promote signal ornamentation. While less common, this alternative outcome offers a unique opportunity to dissect the factors that may lead to different evolutionary pathways. In addition, contrary to traditional assumptions, no sensory modality is completely ‘safe’ as eavesdroppers are ubiquitous and have a broad array of sensory filters that allow opportunity for signal exploitation. We discuss how anthropogenic change affects interactions between eavesdropping enemies and their victims as it rapidly modifies signalling environments and community composition. Drawing on diverse research from a range of taxa and sensory modalities, we synthesize current knowledge on anti-eavesdropper strategies, discuss challenges in this field and highlight fruitful new directions for future research. Ultimately, this review offers a conceptual framework to understand the diverse strategies used by signallers to communicate under the pressure imposed by their eavesdropping enemies.     

Parasitoids, polydnaviruses and endosymbiosis

Symbiotic associations traditionally have been treated as evolutionary curios rather than as a major source of evolutionary innovation. Recent research on a wide variety of organisms is changing this view and is breaking down the barriers between the traditional categories of parasitism, commensalism and mutualism, to produce a more flexible view of multispecific interactions. An especially abundant, but little discussed, mutualism exists between parasitoid wasps in the superfamily Ichneumonoidea and a novel form of DNA viruses known as polydnaviruses. Mutualisms between viruses and eukaryotes are not often reported, although as many as 100 000 species of organisms may exhibit this unusual association. In this review Jim Whitfield considers what is known about the parasitoid-polydnavirus relationship and how (and from what) it might have arisen.     

Key transitions during the evolution of animal phototransduction: novelty, "tree-thinking," co-option, and co-duplication

Biologists are amazed by the intricacy and complexity of biologicalinteractions between molecules, cells, organisms, and ecosystems.Yet underlying all this biodiversity is a universal common ancestry.How does evolution proceed from common starting points to generatethe riotous biodiversity we see today? This "novelty problem"—understandinghow novelty and common ancestry relate—has become of criticalimportance, especially since the realization that genes anddevelopmental processes are often conserved across vast phylogeneticdistances. In particular, two processes have emerged as theprimary generators of diversity in organismal form: duplicationplus divergence and co-option. In this article, we first illustratehow phylogenetic methodology and "tree-thinking" can be usedto distinguish duplication plus divergence from co-option. Second,we review two case studies in photoreceptor evolution—onesuggesting a role for duplication plus divergence, the otherexemplifying how co-option can shape evolutionary change. Finally,we discuss how our tree-thinking approach differs from othertreatments of the origin of novelty that utilized a "linear-thinking"approach in which evolution is viewed as a linear and gradualprogression, often from simple to complex phenotype, drivenby natural selection.     

Evolving views of viral evolution: towards an evolutionary biology of viruses.

Despite considerable interest in viral evolution, at least among virologists, viruses are rarely considered from the same evolutionary vantage point as other organisms. Early work of necessity emphasized phenotype and phenotypic variation (and therefore arguably was more oriented towards the broader biological and ecological perspectives). More recent work (essentially since the development of molecular evolution in the 1960's but beginning earlier) has concentrated on genotypic variation, with less clarity about the significance of such variations. Other aspects of evolutionary theory, especially considerations of natural selection and of evolutionary constraints, have not widely been applied to viruses, and an evolutionary framework for virology has long been lacking. This becomes apparent in considering 'emerging' viruses, which have often been treated on an ad hoc basis. It was often felt that, because previously unrecognized viruses are involved, mechanisms of viral emergence must mirror the unpredictability of mutations in the viral genome. However, most examples of viral emergence are independent of mutation, at least initially, and are often pre-existing viruses in changed circumstances ('viral traffic'). This conclusion also readily follows from ordinary Darwinian premises, which would require that, like other living species, 'new' organisms are descended only from existing species. In this respect, from a Darwinian perspective, viruses would appear to resemble other organisms.     

On mechanistic reasoning in unexpected places: the case of population genetics

A strong case has been made for the role and value of mechanistic reasoning in process-oriented sciences, such as molecular biology and neuroscience. This paper shifts focus to assess the role of mechanistic reasoning in an area where it is neither obvious nor expected: population genetics. Population geneticists abstract away from the causal-mechanical details of individual organisms and, instead, use mathematics to describe population-level, statistical phenomena. This paper, first, develops a framework for the identification of mechanistic reasoning where it is not obvious: mathematical and mechanistic styles of scientific reasoning. Second, it applies this framework to demonstrate that both styles are integrated in modern investigations of evolutionary biology. Characteristic of the former, applied population genetic techniques provide statistical evidence for associations between genotype, phenotype, and fitness. Characteristic of the latter, experimental interventions provide causal-mechanical evidence for associations between the very same relationships, often in the same model organisms. The upshot is a richer perspective of how evolutionary biologists build evidence for hypotheses regarding adaptive evolution and a general framework for assessing the scope of mechanistic reasoning across the sciences.     

Dimetrodon Is Not a Dinosaur: Using Tree Thinking to Understand the Ancient Relatives of Mammals and their Evolution

The line of descent that includes all living mammals extends back in time over 300 million years. Many of the ancient relatives of mammals that fall along this line are very different in appearance from living mammals and are frequently mistaken for reptiles such as dinosaurs. This misconception is reinforced by the fact that these animals are often referred to as “mammal-like reptiles,” a term reflecting outdated methods for classifying organisms. In reality, these ancient mammal-relatives, known as synapsids, are more closely related to living mammals than they are to any reptiles. Evolutionary trees, which depict patterns of descent from common ancestors among organisms, are very useful for understanding why this is the case and for reconstructing the evolutionary histories of many of the unique characters found in mammals. Here, I provide an introduction to evolutionary trees and their implications for understanding the relationships between mammals, synapsids, and reptiles. This is followed by a review of synapsid diversity and a discussion of how evolutionary trees can be used to investigate when in synapsid history different mammalian characteristics first appeared.     

Tsetse--A haven for microorganisms

Arthropods are involved in the transmission of parasitic and viral agents that cause devastating diseases in animals and plants. Effective control strategies for many of these diseases still rely on the elimination or reduction of vector insect populations. In addition to these pathogenic organisms, arthropods are rich in microbes that are symbiotic in their associations and are often necessary for the fecundity and viability of their hosts. Because the viability of the host often depends on these obligate symbionts, and because these organisms often live in close proximity to disease-causing pathogens, they have been of interest to applied biologists as a potential means to genetically manipulate populations of pest species. As knowledge on these symbiotic associations accumulates from distantly related insect taxa, conserved mechanisms for their transmission and evolutionary histories are beginning to emerge. Here, Serap Aksoy summarizes current knowledge on the functional and evolutionary biology of the multiple symbionts harbored in the medically and agriculturally important insect group, tsetse, and their potential role in the control of trypanosomiasis.     

How urbanization affects sexual communication

Urbanization is rapidly altering landscapes worldwide, changing environmental conditions, and creating novel selection pressures for many organisms. Local environmental conditions affect the expression and evolution of sexual signals and mating behaviors; changes in such traits have important evolutionary consequences because of their effect on reproduction. In this review, we synthesize research investigating how sexual communication is affected by the environmental changes associated with urbanization—including pollution from noise, light, and heavy metals, habitat fragmentation, impervious surfaces, urban heat islands, and changes in resources and predation. Urbanization often has negative effects on sexual communication through signal masking, altering condition‐dependent signal expression, and weakening female preferences. Though there are documented instances of seemingly adaptive shifts in trait expression, the ultimate impact on fitness is rarely tested. The field of urban evolution is still relatively young, and most work has tested whether differences occur in response to various aspects of urbanization. There is limited information available about whether these responses represent phenotypic plasticity or genetic changes, and the extent to which observed shifts in sexual communication affect reproductive fitness. Our understanding of how sexual selection operates in novel, urbanized environments would be bolstered by more studies that perform common garden studies and reciprocal transplants, and that simultaneously evaluate multiple environmental factors to tease out causal drivers of observed phenotypic shifts. Urbanization provides a unique testing ground for evolutionary biologists to study the interplay between ecology and sexual selection, and we suggest that more researchers take advantage of these natural experiments. Furthermore, understanding how sexual communication and mating systems differ between cities and rural areas can offer insights on how to mitigate negative, and accentuate positive, consequences of urban expansion on the biota, and provide new opportunities to underscore the relevance of evolutionary biology in the Anthropocene.     

Polar bodies--more a lack of understanding than a lack of respect

Polar bodies are as diverse as the organisms that produce them. Although in many animals these cells often die following meiotic maturation of the oocyte, in other organisms they are an essential and diverse part of embryonic development. Here we highlight some of this diversity and summarize the evolutionary basis for their utility.     

The evolution of condition-dependent sex in the face of high costs

Facultatively sexual organisms often engage in sex more often when in poor condition. We show that such condition-dependent sex carries evolutionary advantages and can explain the evolution of sexual reproduction even when sex entails high costs. Specifically, we show that alleles promoting individuals of low fitness to have sex more often than individuals of high fitness spread through a population. Such alleles are more likely to segregate out of bad genetic backgrounds and onto good genetic backgrounds, where they tend to remain. This "abandon-ship" mechanism provides a plausible model for the evolution and maintenance of facultative sex.     

Adaptive phenotypic plasticity in the Midas cichlid fish pharyngeal jaw and its relevance in adaptive radiation

Background  

Phenotypic evolution and its role in the diversification of organisms is a central topic in evolutionary biology. A neglected factor during the modern evolutionary synthesis, adaptive phenotypic plasticity, more recently attracted the attention of many evolutionary biologists and is now recognized as an important ingredient in both population persistence and diversification. The traits and directions in which an ancestral source population displays phenotypic plasticity might partly determine the trajectories in morphospace, which are accessible for an adaptive radiation, starting from the colonization of a novel environment. In the case of repeated colonizations of similar environments from the same source population this "flexible stem" hypothesis predicts similar phenotypes to arise in repeated subsequent radiations. The Midas Cichlid (Amphilophus spp.) in Nicaragua has radiated in parallel in several crater-lakes seeded by populations originating from the Nicaraguan Great Lakes. Here, we tested phenotypic plasticity in the pharyngeal jaw of Midas Cichlids. The pharyngeal jaw apparatus of cichlids, a second set of jaws functionally decoupled from the oral ones, is known to mediate ecological specialization and often differs strongly between sister-species.     

系统发育研究中“长枝吸引”现象概述

系统发育研究(phylogeny)不仅有助于重建地球所有生物体的进化历史, 而且还可以揭示进化生物学领域中的一些基本问题。清晰了解各生物物种进化历程及不同物种之间的进化关系, 是进一步研究和探索生物学其他学科的基础。但是现今广泛应用的所有系统发育分析方法都存在一定的局限性, 在一定程度上不能有效消除各种误差, 从而不能客观地处理和分析数据, 也就不能成功重建生物进化历程, 真实反映物种进化关系。系统发育研究中, “长枝吸引” (Long-branch Attraction, LBA)假象是最为困扰研究者的问题。文章从“长枝吸引”问题的产生原由、检测方法以及消除策略等多个方面进行详尽概述, 并通过列举典型实例, 阐述了解决“长枝吸引”问题的途径。     

Ecological aspects of the evolutionary processes

Darwin in his On the Origin of species made it clear that evolutionary change depends on the combined action of two different causes, the first being the origin of genetically based phenotypic variation in the individual organisms comprising the population and the second being the action of selective agents of the external environment placing demands on the individual organisms. For over a century following Darwin, most evolutionists focused on the origin of inherited variation and its transmission; many workers continue to regard genetics to be the core of evolutionary theory. Far less attention has been given to the exact nature of the selective agents with most evolutionists still treating this cause imprecisely to the detriment of our understanding of both nomological and historical evolutionary theory. Darwin was vague in the meaning of his new concept of "Natural Selection," using it interchangeably as one of the causes for evolutionary change and as the final outcome (= evolutionary change). In 1930, natural selection was defined clearly as "non-random, differential reproduction of genes" by R. Fisher and J.B.S. Haldane which is a statement of the outcome of evolutionary process and which omits mention of the causes bringing about this change. Evolutionists quickly accepted this outcome definition of natural selection, and have used interchangeably selection both as a cause and as the result of evolutionary change, causing great confusion. Herein, the details will be discussed of how the external environment (i.e., the environment-phenotype interaction) serves as selective agents and exerts demands on the phenotypic organisms. Included are the concepts of fitness and of the components of fitness (= adaptations) which are respectively (a) survival, (b) direct reproductive and (c) indirect reproductive features. Finally, it will be argued that historical-narrative analyses of organisms, including classification and phylogenetic history, are possible only with a full understanding of nomological evolutionary theory and with functional/adaptive studies of the employed taxonomic features in addition to the standard comparative investigations.     

Cooperation and conflict in the evolution of individuality. IV. Conflict mediation and evolvability in Volvox carteri

The continued well being of evolutionary individuals (units of selection and evolution) depends upon their evolvability, that is their capacity to generate and evolve adaptations at their level of organization, as well as their longer term capacity for diversifying into more complex evolutionary forms. During a transition from a lower- to higher-level individual, such as the transition between unicellular and multicellular organisms, the evolvability of the lower-level (cells) must be restricted, while the evolvability of the new higher-level unit (multicellular organism) must be enhanced. For these reasons, understanding the factors leading to an evolutionary transition should help us to understand the factors underlying the emergence of evolvability of a new evolutionary unit. Cooperation among lower-level units is fundamental to the origin of new functions in the higher-level unit. Cooperation can produce a new more complex evolutionary unit, with the requisite properties of heritable fitness variations, because cooperation trades fitness from a lower-level (the costs of cooperation) to the higher-level (the benefits for the group). For this reason, the evolution of cooperative interactions helps us to understand the origin of new and higher-levels of fitness and organization. As cooperation creates a new level of fitness, it also creates the opportunity for conflict between levels of selection, as deleterious mutants with differing effects at the two levels arise and spread. This conflict can interfere with the evolvability of the higher-level unit, since the lower and higher-levels of selection will often "disagree" on what adaptations are most beneficial to their respective interests. Mediation of this conflict is essential to the emergence of the new evolutionary unit and to its continued evolvability. As an example, we consider the transition from unicellular to multicellular organisms and study the evolution of an early-sequestered germ-line in terms of its role in mediating conflict between the two levels of selection, the cell and the cell group. We apply our theoretical framework to the evolution of germ/soma differentiation in the green algal group Volvocales. In the most complex member of the group, Volvox carteri, the potential conflicts among lower-level cells as to the "right" to reproduce the higher-level individual (i.e. the colony) have been mediated by restricting immortality and totipotency to the germ-line. However, this mediation, and the evolution of an early segregated germ-line, was achieved by suppressing mitotic and differentiation capabilities in all post-embryonic cells. By handicapping the soma in this way, individuality is ensured, but the solution has affected the long-term evolvability of this lineage. We think that although conflict mediation is pivotal to the emergence of individuality at the higher-level, the way in which the mediation is achieved can greatly affect the longer-term evolvability of the lineage.     

ANDROGENETICS AND TRIPLOIDS FROM AN INTERACTING PARTHENOGENETIC HYBRID AND ITS ANCESTORS IN STICK INSECTS

Populations of unisexual organisms are often assumed to be genetically invariant (clones) and destined to a short existence on an evolutionary timescale. Unisexual organisms are most often obligate parthenogens and, by definition, ought to be completely isolated reproductively from related bisexual organisms. The assumption of complete reproductive isolation between amphimictic ancestors and thelytokous hybrids is common to most hypotheses on the evolution of sex and its adaptive significance. Stick insects of the genus Bacillus however provide evidence for reproductive interactions between allodiploid parthenogens and their ancestors, because pure species progeny (androgenetics) and triploid descendants are produced. These findings demonstrate that, through androgenesis, offspring of parthenogenetic hybrid females can contribute specimens of both sexes to the fathering species when fertilized by syntopic ancestral males and the parthenogenetic egg of strictly clonal females, when fertilized, allows a third genome to be added to the allodiploid chromosome set. These triploid genomes promote further genetic diversification and evolution of the unisexual populations through the formation of new clones by recombination during the changed maturation mode of allotriploid eggs. All this argues for much more complex breeding systems and evolutionary pathways than are usually assumed for hybrid unisexual organisms.