Genetics,development and composition of the insect head – A beetle’s view

Many questions regarding evolution and ontogeny of the insect head remain open. Likewise, the genetic basis of insect head development is poorly understood. Recently, the investigation of gene expression data and the analysis of patterning gene function have revived interest in insect head development. Here, we argue that the red flour beetle Tribolium castaneum is a well suited model organism to spearhead research with respect to the genetic control of insect head development. We review recent molecular data and discuss its bearing on early development and morphogenesis of the head. We present a novel hypothesis on the ontogenetic origin of insect head sutures and review recent insights into the question on the origin of the labrum. Further, we argue that the study of developmental genes may identify the elusive anterior non-segmental region and present some evidence in favor of its existence. With respect to the question of evolution of patterning we show that the head Anlagen of the fruit fly Drosophila melanogaster and Tribolium differ considerably and we review profound differences of their genetic regulation. Finally, we discuss which insect model species might help us to answer the open questions concerning the genetic regulation of head development and its evolution.     

The evolution of HIV: inferences using phylogenetics

Molecular phylogenetics has revolutionized the study of not only evolution but also disparate fields such as genomics, bioinformatics, epidemiology, ecology, microbiology, molecular biology and biochemistry. Particularly significant are its achievements in population genetics as a result of the development of coalescent theory, which have contributed to more accurate model-based parameter estimation and explicit hypothesis testing. The study of the evolution of many microorganisms, and HIV in particular, have benefited from these new methodologies. HIV is well suited for such sophisticated population analyses because of its large population sizes, short generation times, high substitution rates and relatively small genomes. All these factors make HIV an ideal and fascinating model to study molecular evolution in real time. Here we review the significant advances made in HIV evolution through the application of phylogenetic approaches. We first examine the relative roles of mutation and recombination on the molecular evolution of HIV and its adaptive response to drug therapy and tissue allocation. We then review some of the fundamental questions in HIV evolution in relation to its origin and diversification and describe some of the insights gained using phylogenies. Finally, we show how phylogenetic analysis has advanced our knowledge of HIV dynamics (i.e., phylodynamics).     

Slit/Robo-mediated axon guidance in Tribolium and Drosophila: divergent genetic programs build insect nervous systems

As the complexity of animal nervous systems has increased during evolution, developmental control of neuronal connectivity has become increasingly refined. How has functional diversification within related axon guidance molecules contributed to the evolution of nervous systems? To address this question, we explore the evolution of functional diversity within the Roundabout (Robo) family of axon guidance receptors. In Drosophila, Robo and Robo2 promote midline repulsion, while Robo2 and Robo3 specify the position of longitudinal axon pathways. The Robo family has expanded by gene duplication in insects; robo2 and robo3 exist as distinct genes only within dipterans, while other insects, like the flour beetle Tribolium castaneum, retain an ancestral robo2/3 gene. Both Robos from Tribolium can mediate midline repulsion in Drosophila, but unlike the fly Robos cannot be down-regulated by Commissureless. The overall architecture and arrangement of longitudinal pathways are remarkably conserved in Tribolium, despite it having only two Robos. Loss of TcSlit causes midline collapse of axons in the beetle, a phenotype recapitulated by simultaneous knockdown of both Robos. Single gene knockdowns reveal that beetle Robos have specialized axon guidance functions: TcRobo is dedicated to midline repulsion, while TcRobo2/3 also regulates longitudinal pathway formation. TcRobo2/3 knockdown reproduces aspects of both Drosophila robo2 and robo3 mutants, suggesting that TcRobo2/3 has two functions that in Drosophila are divided between Robo2 and Robo3. The ability of Tribolium to organize longitudinal axons into three discrete medial–lateral zones with only two Robo receptors demonstrates that beetle and fly achieve equivalent developmental outcomes using divergent genetic programs.     

A mathematical model for animal ecology with special reference to the ecology of the Tribolium beetle

An axiomatic model, originally developed for evolutionary theory, is adapted to describe animal ecology. A matrix representation of the present simulation of Tribolium castaneum, genetic strain c-IVa, is incorporated in the general system. The “weal” of a population is introduced as part of the general system and specified in more detail for Tribolium. The application of the model to the principle of competitive displacement is discussed.     

Genome-wide acceleration of protein evolution in flies (Diptera)

Background

The rate of molecular evolution varies widely between proteins, both within and among lineages. To what extent is this variation influenced by genome-wide, lineage-specific effects? To answer this question, we assess the rate variation between insect lineages for a large number of orthologous genes.

Results

When compared to the beetle Tribolium castaneum, we find that the stem lineage of flies and mosquitoes (Diptera) has experienced on average a 3-fold increase in the rate of evolution. Pairwise gene comparisons between Drosophila and Tribolium show a high correlation between evolutionary rates of orthologous proteins.

Conclusion

Gene specific divergence rates remain roughly constant over long evolutionary times, modulated by genome-wide, lineage-specific effects. Among the insects analysed so far, it appears that the Tribolium genes show the lowest rates of divergence. This has the practical consequence that homology searches for human genes yield significantly better matches in Tribolium than in Drosophila. We therefore suggest that Tribolium is better suited for comparisons between phyla than the widely employed dipterans.     

甲虫群落构建研究进展

甲虫作为节肢动物中一个庞大的类群,具有繁简各异的生活习性,在生态系统中发挥极为重要的作用.甲虫群落构建是群落生态研究的重要内容之一,可揭示不同生态环境下生物多样性的形成与维持机制.现阶段对甲虫群落构建的研究主要通过物种的功能性状,或将系统发育结构与物种功能性状相结合的方法,对群落构建过程进行解析,用以判断主要的驱动因素...     

Freezing or death feigning? Beetles selected for long death feigning showed different tactics against different predators

Prey evolve antipredator strategies against multiple enemies in nature. We examined how a prey species adopts different predation avoidance tactics against pursuit or sit‐and‐wait predators. As prey, we used three strains of Tribolium beetles artificially selected for short (short strain) or long (long strain) duration of death feigning, and a stock culture (base population). Death feigning is known to be effective for evading a jumping spider in the case of the long strains, while the present study showed that the long‐strain beetles used freezing against a sit‐and‐wait type predator, Amphibolus venator, in this study. The short‐ strain beetles were more easily oriented toward predators. The time to predation was also shorter in the short strains compared to the long strains. The results showed that, as prey, the short strains displayed the same behavior, escaping, against both types of predators. Traditionally, death feigning has been thought to be the last resort in a series of antipredator avoidance behaviors. However, our results showed that freezing and death feigning were not parts of a series of behaviors, but independent strategies against different predators, at least for long‐strain beetles. We also examined the relationship between a predator''s starvation level and its predatory behavior. In addition, the orientation behavior toward and predation rate on the prey were observed to determine how often the predatory insect attacked the beetles.     

Entomopathogenic Nematodes as a Model System for Advancing the Frontiers of Ecology

Entomopathogenic nematodes (EPNs) in the families Heterorhabditidae and Steinernematidae have a mutualistic–symbiotic association with enteric γ-Proteobacteria (Steinernema–Xenorhabdus and Heterorhabditis–Photorhabdus), which confer high virulence against insects. EPNs have been studied intensively because of their role as a natural mortality factor for soil-dwelling arthropods and their potential as biological control agents for belowground insect pests. For many decades, research on EPNs focused on the taxonomy, phylogeny, biogeography, genetics, physiology, biochemistry and ecology, as well as commercial production and application technologies. More recently, EPNs and their bacterial symbionts are being viewed as a model system for advancing research in other disciplines such as soil ecology, symbiosis and evolutionary biology. Integration of existing information, particularly the accumulating information on their biology, into increasingly detailed population models is critical to improving our ability to exploit and manage EPNs as a biological control agent and to understand ecological processes in a changing world. Here, we summarize some recent advances in phylogeny, systematics, biogeography, community ecology and population dynamics models of EPNs, and describe how this research is advancing frontiers in ecology.     

The contribution of evening primrose (<Emphasis Type="Italic">Oenothera biennis</Emphasis>) to a modern synthesis of evolutionary ecology

In this review, I consider the contribution that common evening primrose (Oenothera biennis) has made towards integrating the ecology, evolution and genetics of species interactions. Oenothera biennis was among the earliest plant models in genetics and cytogenetics and it played an important role in the modern synthesis of evolutionary biology. More recently, population and ecological genetics approaches have provided insight into the patterns of genetic variation within and between populations, and how a combination of abiotic and biotic factors maintain and select on heritable variation within O. biennis populations. From an ecological perspective, field experiments show that genetic variation and evolution within populations can have cascading effects throughout communities. Plant genotype affects the preference and performance of individual arthropod populations, as well as the composition, biomass, total abundance and diversity of arthropod species on plants. A combination of experiments and simulation models show that natural selection on specific plant traits can drive rapid ecological changes in these same community variables. At the patch level, increasing genotypic diversity leads to a greater abundance and diversity of omnivorous and predaceous arthropods, which is also associated with increased biomass and fecundity of plants in genetically diverse patches. Finally, in questioning whether a community genetics perspective is needed in biology, I review several multifactorial experiments which show that plant genotype often explains as much variation in community variables as other ecological factors typically identified as most important in ecology. As a whole, research in the O. biennis system has contributed to a more complete understanding of the dynamic interplay between ecology, evolution and genetics.     

Bridging the gap between theory and data in ecological models

In September 2011, the University of Essex, UK, hosted an interdisciplinary conference, Mathematical and Theoretical Ecology 2011 (MATE 2011), with the theme of ‘Linking models with ecological processes’. The aim of the meeting was to create discussion and debate between modellers and empiricists working in ecology. A wide range of topics were discussed at the meeting including evolutionary and community models of ecosystem structure, epidemiological models, non-linear models of population dynamics, spatiotemporal models, individual and collective movement behaviour, and applications of ecological models to engineering problems. In this introductory article, we provide a report of the MATE 2011 meeting, and briefly review the most recent relevant research in the fields of mathematical and theoretical ecology. We introduce and summarise the eight contributed articles that were selected for this special issue. The diverse range of topics and the wide range of mathematical, statistical and computational tools used illustrate the broad appeal and depth of research in the rich field of mathematical and theoretical ecology.     

The integrative biology of genetic dominance

Dominance is a basic property of inheritance systems describing the link between a diploid genotype at a single locus and the resulting phenotype. Models for the evolution of dominance have long been framed as an opposition between the irreconcilable views of Fisher in 1928 supporting the role of largely elusive dominance modifiers and Wright in 1929, who viewed dominance as an emerging property of the structure of enzymatic pathways. Recent theoretical and empirical advances however suggest that these opposing views can be reconciled, notably using models investigating the regulation of gene expression and developmental processes. In this more comprehensive framework, phenotypic dominance emerges from departures from linearity between any levels of integration in the genotype-to-phenotype map. Here, we review how these different models illuminate the emergence and evolution of dominance. We then detail recent empirical studies shedding new light on the diversity of molecular and physiological mechanisms underlying dominance and its evolution. By reconciling population genetics and functional biology, we hope our review will facilitate cross-talk among research fields in the integrative study of dominance evolution.     

Reproductive interference in laboratory experiments of interspecific competition

Many studies that have researched interspecific competition in Callosobruchus (bean beetles), Drosophila (fruit flies), and Tribolium (flour beetles) have considered the major drivers of interspecific competition to be interspecific resource competition and intraguild cannibalism. These competition drivers have a density-dependent effect on the population dynamics. However, some studies have also detected a relative-frequency-dependent effect in the observed population dynamics. The most likely causal mechanism of this relative frequency dependence is reproductive interference, defined as any interspecific sexual interaction that damages female reproductive success. Reproductive interference has been overlooked by most laboratory studies in spite of the critical effect on the competition outcome. In this paper, I review laboratory studies of these insect genera from the perspective of reproductive interference and show that the reported results can be more reasonably interpreted by the joint action of reproductive interference and resource competition, including intraguild cannibalism. In addition, on the basis of results reported by a small number of related studies, I discuss the behavioral and evolutionary changes induced in those genera by reproductive interference.     

An integrative approach to understanding host–parasitoid population dynamics in real landscapes

Many of our advances regarding the spatial ecology of predators and prey have been attributed to research with insect parasitoids and their hosts. Host–parasitoid systems are ideal for spatial-ecological studies because of the small size of the organisms, the often discrete distribution of their resources, and the relative ease with which host mortality from parasitoids can be determined. We outline an integrated approach to studying host–parasitoid interactions in heterogeneous natural landscapes. This approach involves conducting experiments to obtain critically important information on dispersal and boundary behavior of the host and parasitoid, large-scale manipulations of landscape structure to reveal the impacts of landscape change on host–parasitoid interactions and temporal population dynamics, and the development of spatially realistic, behavior-based landscape models. The dividends from such an integrative approach are far reaching, as is illustrated in our research on the prairie planthopper Prokelisia crocea and its egg parasitoid Anagrus columbi that occurs in the tall-grass prairies of North America. Here, we describe the population structure of this system which is based on a long-term survey of planthoppers and parasitoids among host–plant patches. We also outline novel approaches to experimentally quantify and model the movement and boundary behavior of animals in general. The value of this information is revealed in a landscape-level field experiment that was designed to test predictions about how landscape change affects the spatial and temporal population dynamics of the host and parasitoid. Finally, with these empirical data as the foundation, we describe novel simulation models that are spatially realistic and behavior based. Drawing from this integrated approach and case study, we identify key research questions for the future.     

Dispersal and reproduction in Tribolium: The influence of food level

Cohorts of 100 newly eclosed adults of strains bI and bIV of T. confusum and strains cI and cIV of T. castaneum emigrated over a 15-day interval from experimental habitats which contained a constant volume of 8, 4, 1, and 0 g of flour diluted with powdered polyethylene. Food level affected the emigration of T. confusum adults only during the interval before the normal onset of reproduction. Thereafter the beetles exhibited virtually identical patterns of departure. Larval recruitment did not decline when food levels ranged from 8 to 1. T. castaneum delayed their emigration at progressively lower food levels, but also produced fewer progeny. In strain cI, where emigration was more strongly deferred, the decline in larval recruitment was less pronounced. Both species of Tribolium departed most rapidly when starved outright, but only after an initial delay.The results are interpreted from the point of view of the oögenesis-flight syndrome. It is suggested that the Tribolium system provides an opportunity to relate differences in colonizing ability among several closely related species to specific differences in the physiological mechanism of adaptive dispersal.     

Analysis of orientation behaviour of Tribolium castaneum and T. confusum to synthetic aggregation pheromone

Adult male T. castaneum (Herbst) and T. confusum (du Val) secretes an aggregation pheromone that is attractive to both sexes. Orientation behaviour of the two Tribolium species responding to different concentrations of synthetic aggregation pheromone in still and moving air was studied in an 2.5 m×0.4 m olfactometer. Analysis of Tribolium tracks indicated that the aggregation pheromone stimulated the beetles to walk faster at higher concentrations to increase the frequency and magnitude of turning and to decrease track reversal distances and distances between turns. The mean walking speed of both species was lowest at the highest air speed. The behavioural responses of the beetles to the pheromone in still and moving air were similar, indicating chemotaxis as the major orientation mechanism used by both species to locate an odour source. The beetles showed greater orientation efficiency within a discrete pheromone plume than a diffuse plume.     

Exploring systemic RNA interference in insects: a genome-wide survey for RNAi genes in Tribolium

Background

RNA interference (RNAi) is a highly conserved cellular mechanism. In some organisms, such as Caenorhabditis elegans, the RNAi response can be transmitted systemically. Some insects also exhibit a systemic RNAi response. However, Drosophila, the leading insect model organism, does not show a robust systemic RNAi response, necessitating another model system to study the molecular mechanism of systemic RNAi in insects.

Results

We used Tribolium, which exhibits robust systemic RNAi, as an alternative model system. We have identified the core RNAi genes, as well as genes potentially involved in systemic RNAi, from the Tribolium genome. Both phylogenetic and functional analyses suggest that Tribolium has a somewhat larger inventory of core component genes than Drosophila, perhaps allowing a more sensitive response to double-stranded RNA (dsRNA). We also identified three Tribolium homologs of C. elegans sid-1, which encodes a possible dsRNA channel. However, detailed sequence analysis has revealed that these Tribolium homologs share more identity with another C. elegans gene, tag-130. We analyzed tag-130 mutants, and found that this gene does not have a function in systemic RNAi in C. elegans. Likewise, the Tribolium sid-like genes do not seem to be required for systemic RNAi. These results suggest that insect sid-1-like genes have a different function than dsRNA uptake. Moreover, Tribolium lacks homologs of several genes important for RNAi in C. elegans.

Conclusion

Although both Tribolium and C. elegans show a robust systemic RNAi response, our genome-wide survey reveals significant differences between the RNAi mechanisms of these organisms. Thus, insects may use an alternative mechanism for the systemic RNAi response. Understanding this process would assist with rendering other insects amenable to systemic RNAi, and may influence pest control approaches.     

Evolutionary divergence of the paralogs Methoprene tolerant (Met) and germ cell expressed (gce) within the genus Drosophila

Juvenile hormone (JH) signaling underpins both regulatory and developmental pathways in insects. However, the JH receptor is poorly understood. Methoprene tolerant (Met) and germ cell expressed (gce) have been implicated in JH signaling in Drosophila. We investigated the evolution of Met and gce across 12 Drosophila species and found that these paralogs are conserved across at least 63 million years of dipteran evolution. Distinct patterns of selection found using estimates of dN/dS ratios across Drosophila Met and gce coding sequences, along with their incongruent temporal expression profiles in embryonic Drosophila melanogaster, illustrate avenues through which these genes have diverged within the Diptera. Additionally, we demonstrate that the annotated gene CG15032 is the 5′ terminus of gce.In mosquitoes and beetles, a single Met-like homolog displays structural similarity to both Met and gce, and the intron locations are conserved with those of gce. We found that Tribolium and mosquito Met orthologs are assembled from Met- and gce-specific domains in a modular fashion. Our results suggest that Drosophila Met and gce experienced divergent evolutionary pressures following the duplication of an ancestral gce-like gene found in less derived holometabolous insects.     

Linking biodiversity and ecosystems: towards a unifying ecological theory

Community ecology and ecosystem ecology provide two perspectives on complex ecological systems that have largely complementary strengths and weaknesses. Merging the two perspectives is necessary both to ensure continued scientific progress and to provide society with the scientific means to face growing environmental challenges. Recent research on biodiversity and ecosystem functioning has contributed to this goal in several ways. By addressing a new question of high relevance for both science and society, by challenging existing paradigms, by tightly linking theory and experiments, by building scientific consensus beyond differences in opinion, by integrating fragmented disciplines and research fields, by connecting itself to other disciplines and management issues, it has helped transform ecology not only in content, but also in form. Creating a genuine evolutionary ecosystem ecology that links the evolution of species traits at the individual level, the dynamics of species interactions, and the overall functioning of ecosystems would give new impetus to this much-needed process of unification across ecological disciplines. Recent community evolution models are a promising step in that direction.     

Application of flow cytometry for exploring the evolution of Geosmithia fungi living in association with bark beetles: the role of conidial DNA content

Geosmithia belongs among fungi living in symbiosis with phloem-feeding bark beetles. Several species have altered their ecology to that of obligatory symbiosis with ambrosia beetles, which has led to a shift in their phenotype and caused formation of large spherical conidia. In this study, we pose the following questions; (1) Is the conidial DNA content of Geosmithia correlated with conidial volume?; (2) Is the DNA content of Geosmithia related to the degree of mutual dependence between Geosmithia and their vector? There was a positive and strong correlation between conidial DNA content and conidial volume in Geosmithia. Also species more narrowly associated with the vector tend to have a larger conidial DNA content and volume than less narrowly associated species. Ambrosia fungi achieved the biggest conidial DNA content and volume compared to other species. We suppose that polyploidisation occurred during the evolution of ambrosia species in the genus Geosmithia.     

The effects of temperature,relative humidity,light, and resource quality on flight initiation in the red flour beetle,Tribolium castaneum

We investigated the environmental conditions that induce a flight response in the red flour beetle, Tribolium castaneumHerbst (Coleoptera: Tenebrionidae), including resource quality, temperature, relative humidity, and light. Over 72‐h trial periods, we observed the proportion of individuals emigrating by flight to range from 0.0 in extreme heat or cold to 0.82 with starvation. Resource quality, presence of a light source, and temperature all directly influenced the initiation of the flight response. We did not detect any effect of relative humidity or sudden change in temperature on the incidence of flight. We discuss our findings in the context of Tribolium ecology and evolution.