Adaptive Radiation in Insects and Plants: Time and Opportunity

SYNOPSIS. Insects and their hostplants represent the major partof terrestrial diversity, yet we are just beginning to understandwhy there are so very many species. By far the most influentialmodel of insect/plant diversification has been Ehrlich and Raven's(1964) hypothesis of insect/plant coevolution. While the coevolutionmodel was based on macroevolutionary patterns in plant defensesand hostplant affiliations, most of the subsequent work hasbeen on its possible ecological and genetic mechanisms, withrelatively little systematic scrutiny of the evolutionary patternsEhrlich and Raven described. We explore the possible roles insect/plantinteractions may play in the long-term evolution of insect andplant lineages, and review some of the evidence on whether ornot insects and plants have exerted reciprocal influences oneach other's diversification. Insects and plants have diversified over roughly the same timeintervals, and many insect host/affiliations are evolutionarilyconserved, thus reflecting long/term, phylogenetic history.Rather than accumulating herbivores at a rate proportional totheir geographic area of distribution or biomass, some plantgroups pose apparent chemical barriers to potential herbivorecolonists, and seem accessible to relatively few insect lineages,possibly preadapted by use of chemically similar or relatedhostplants. Evolutionary innovations in plant defenses and ininsect feeding habits seem to have spurred their respectiveadaptive radiations, thus ecological opportunity may influencelong-term evolutionary success. The greater diversity of insectsand plants in the tropics, compared to the temperate zone, probablyreflects the greater age of tropical habitats as well as climaticbarriers that limit successful invasion of the temperate zoneto just those primitively tropical groups able to evolve strategiesfor both overwintering and use of temperate resources. Thoughevidence is still sparse, successful invasion of the temperatezone may promote subsequent radiations of both insects and plants. We conclude that much of the available evidence from systematicsis consistent with Ehrlich and Raven's suggestion that muchof insect and plant diversification has been spurred by a seriesof ecological opportunities over evolutionary time.     

On the move: sloths and their epibionts as model mobile ecosystems

Sloths are unusual mobile ecosystems, containing a high diversity of epibionts living and growing in their fur as they climb slowly through the canopies of tropical forests. These epibionts include poorly studied algae, arthropods, fungi, and bacteria, making sloths likely reservoirs of unexplored biodiversity. This review aims to identify gaps and eliminate misconceptions in our knowledge of sloths and their epibionts, and to identify key questions to stimulate future research into the functions and roles of sloths within a broader ecological and evolutionary context. This review also seeks to position the sloth fur ecosystem as a model for addressing fundamental questions in metacommunity and movement ecology. The conceptual and evidence-based foundation of this review aims to serve as a guide for future hypothesis-driven research into sloths, their microbiota, sloth health and conservation, and the coevolution of symbioses in general.     

Phloem sap proteins: their identities and potential roles in the interaction between plants and phloem-feeding insects

The phloem is a well-known target of sucking and piercing insects that utilize the transported fluid as their major nutrient source. In addition to small molecules like sugars and amino acids, phloem sap of higher land plants contains proteins that can accumulate up to high concentrations. Although the knowledge about the identities of these phloem sap proteins is increasing, the functions of most of them are still poorly understood. Since many phloem sap proteins have predicted roles in wound and defence responses, they constitute a class of compounds that can potentially influence plant-insect interactions. However, there are as yet no studies published that have examined direct effects of phloem sap proteins on insect feeding or vice versa. This review summarizes the current knowledge about the identities of phloem sap proteins, focused on polypeptides with probable functions in wound and defence reactions, and their potential impact on plant-insect interactions is discussed.     

昆虫共生微生物在病虫害和疾病控制上的应用前景

昆虫与微生物之间的互利共生关系是自然界中一种常见的互作形式。昆虫的种类丰富多样并且在自然界中分布广泛,在一定程度上得益于共生微生物的帮助。随着生物技术的不断发展,越来越多的共生微生物和互利共生模式得以发现并深入研究。微生物不仅能够为昆虫的生长发育提供营养,还能合成很多生物活性物质、调节宿主的免疫、对抗捕食者和抵御病原微生物感染,成为宿主昆虫健康和适应的守护者。鉴于共生微生物与昆虫生理生态的密切联系,以及昆虫对人类经济与健康的重要影响,利用共生微生物对昆虫及虫媒病进行生物控制已经成为一个热点研究方向,并展现了良好的应用前景。本文对昆虫共生微生物的多样性、生物学功能、与宿主相互作用机制及其在病虫害和虫媒病防治中的研究进展进行综述和展望。     

Plant scent and plant–insect interactions—Review and outlook from a macroevolutionary perspective

The astonishing diversity of plants and insects and their entangled interactions are cornerstones in terrestrial ecosystems. Co-occurring with species diversity is the diversity of plant secondary metabolites (PSMs). So far, their estimated number is more than 200 000 compounds, which are not directly involved in plant growth and development but play important roles in helping plants handle their environment including the mediation of plant–insect interactions. Here, we use plant volatile organic compounds (VOCs), a key olfactory communication channel that mediates plant–insect interactions, as a showcase of PSMs. In spite of the cumulative knowledge of the functional, ecological, and microevolutionary roles of VOCs, we still lack a macroevolutionary understanding of how they evolved with plant–insect interactions and contributed to species diversity throughout the long coevolutionary history of plants and insects. We first review the literature to summarize the current state-of-the-art research on this topic. We then present various relevant types of phylogenetic methods suitable to answer macroevolutionary questions on plant VOCs and suggest future directions for employing phylogenetic approaches in studying plant VOCs and plant–insect interactions. Overall, we found that current studies in this field are still very limited in their macroevolutionary perspective. Nevertheless, with the fast-growing development of metabolome analysis techniques and phylogenetic methods, it is becoming increasingly feasible to integrate the advances of these two areas. We highlight promising approaches to generate new testable hypotheses and gain a mechanistic understanding of the macroevolutionary roles of chemical communication in plant–insect interactions.     

Parasitoid polydnaviruses: evolution,pathology and applications

One of the more unusual groups of insect pathogens consists of members of the family Polydnaviridae, insect DNA viruses that live in mutual symbioses with their associated parasitoid wasp (Hymentoptera) carriers until they are injected into specific lepidopteran hosts. Once inside this secondary host, polydnaviruses cause a wide variety of negative effects that ultimately ensure the survival of the parasitoid larvae. Because of their unusual life strategy and genetic features, it had been difficult to fully characterise polydnaviruses in terms of evolutionary history, replication cycle and functions in the host that might normally be well characterised for more conventional viruses. Recently, our understanding of polydnavirus evolutionary origins, gene content, genome organisation and functions in parasitism has greatly increased. Key findings are summarised in this review with emphasis on evolution of polydnavirus genes and genomes, their functional roles in insect pathology and their potential applications in insect biological control and biotechnology.     

植物病原-媒介昆虫互作:研究进展与展望

大多数植物病毒及一些植物病原细菌由介体昆虫传播。植物病原与介体昆虫关系的研究有助于找到防控介体传播病原的关键环节,因此植物病原与介体昆虫的互作关系是植物病原传播机理研究中的核心问题。本文概述了国内外在植物病原与介体昆虫互作研究的最新进展,推介了本专辑论文的主要内容,并在此基础上,从生态和进化的角度提出了在植物病原-媒介昆虫互作研究中以下3个值得关注的研究方向:(1)植物病原与介体昆虫互作对生态系统的影响;(2)昆虫介体传播植物病毒的不同方式之间的关联性以及病毒、介体和植物之间的协同进化关系;(3)自然条件下植物病原-媒介昆虫互作的机理。植物病原与媒介昆虫互作的研究,既是生态和进化的理论问题,也和植物病原及其介体昆虫的绿色防控密切相关。     

Evolutionary dynamics of specialisation in herbivorous stick insects

Understanding the evolutionary dynamics underlying herbivorous insect mega‐diversity requires investigating the ability of insects to shift and adapt to different host plants. Feeding experiments with nine related stick insect species revealed that insects retain the ability to use ancestral host plants after shifting to novel hosts, with host plant shifts generating fundamental feeding niche expansions. These expansions were, however, not accompanied by expansions of the realised feeding niches, as species on novel hosts are generally ecologically specialised. For shifts from angiosperm to chemically challenging conifer hosts, generalist fundamental feeding niches even evolved jointly with strong host plant specialisation, indicating that host plant specialisation is not driven by constraints imposed by plant chemistry. By coupling analyses of plant chemical compounds, fundamental and ecological feeding niches in multiple insect species, we provide novel insights into the evolutionary dynamics of host range expansion and contraction in herbivorous insects.     

Integrating ancient patterns and current dynamics of insect–plant interactions: Taxonomic and geographic variation in herbivore specialization

Abstract The search for pattern in the ecology and evolutionary biology of insect–plant associations has fascinated biologists for centuries. High levels of tropical (low-latitude) plant and insect diversity relative to poleward latitudes and the disproportionate abundance of host-specialized insect herbivores have been noted. This review addresses several aspects of local insect specialization, host use abilities (and loss of these abilities with specialization), host-associated evolutionary divergence, and ecological (including “hybrid”) speciation, with special reference to the generation of biodiversity and the geographic and taxonomic identification of “species borders” for swallowtail butterflies (Papilionidae). From ancient phytochemically defined angiosperm affiliations that trace back millions of years to recent and very local specialized populations, the Papilionidae (swallowtail butterflies) have provided a model for enhanced understanding of localized ecological patterns and genetically based evolutionary processes. They have served as a useful group for evaluating the feeding specialization/physiological efficiency hypothesis. They have shown how the abiotic (thermal) environment interacts with host nutrirional suitability to generate “voltinism/suitability” gradients in specialization or preference latitudinally, and geographical mosaics locally. Several studies reviewed here suggest strongly that the oscillation hypothesis for speciation does have considerable merit, but at the same time, some species-level host specializations may lead to evolutionary dead-ends, especially with rapid environmental/habitat changes involving their host plants. Latitudinal gradients in species richness and degree of herbivore feeding specialization have been impacted by recent developments in ecological genetics and evolutionary ecology. Localized insect–plant associations that span the biospectrum from polyphenisms, polymorphisms, biotypes, demes, host races, to cryptic species, remain academically contentious, with simple definitions still debated. However, molecular analyses combined with ecological, ethological and physiological studies, have already begun to unveil some answers for many important ecological/evolutionary questions.     

Community heterogeneity and the evolution of interactions between plants and insect herbivores

Plant communities vary tremendously in terms of productivity, species diversity, and genetic diversity within species. This vegetation heterogeneity can impact both the likelihood and strength of interactions between plants and insect herbivores. Because altering plant-herbivore interactions will likely impact the fitness of both partners, these ecological effects also have evolutionary consequences. We review several hypothesized and well-documented mechanisms whereby variation in the plant community alters the plant-herbivore interaction, discuss potential evolutionary outcomes of each of these ecological effects, and conclude by highlighting several avenues for future research. The underlying theme of this review is that the neighborhood of plants is an important determinant of insect attack, and this results in feedback effects on the plant community. Because plants exert selection on herbivore traits and, reciprocally, herbivores exert selection on plant-defense traits, variation in the plant community likely contributes to spatial and temporal variation in both plant and insect traits, which could influence macroevolutionary patterns.     

Biological functions of carotenoids--diversity and evolution

Carotenoids first emerged in archaebacteria as lipids reinforcing cell membranes. To serve this function their long molecules have extremely rigid backbone due to the linear chain of usually 10 to 11 conjugated C=C bonds in transconfiguration--the length corresponding the thickness of hydrophobic zone of membrane which they penetrate as "molecular rivets". Carotenoids retain their membrane-reinforcing function in some fungi and animals. The general structure of carotenoid molecule, originally having evolved for mechanical functions in membranes, possess a number of other properties that were later used for independent functions. The most striking fact is that these properties proved to fit some new functions to perfection. The polyene chain of 9-11 double bonds absorbs light precisely in the gap of chlorophyll absorption--function as accessory light-harvesting pigments in all plants; Unique arrangement of electronic levels owing to the by polyene chain structure makes carotenoids the only natural compounds capable of excitation energy transfer both (i) from carotenoid excited state to chlorophyll in the light-harvesting complex and (ii) from triplet chlorophyll or singlet oxygen to carotenoid in photosynthetic reaction centers--protection of RC from photodamage. The linear system of conjugated C=C bonds provides high reducing potential of carotenoid molecules making them potent antioxidants in lipid formations. Still, there is a lack of evidence of the chemical antioxidant function of carotenoids, especially in higher organisms; most data demonstrate an antioxidant ability rather than a function. Carotenoids have many other independent biological functions, including: specific coloration patterns in plants and animals, screening from excessive light and spectral filtering, defense of egg proteins from proteases in some invertebrates; the direct carotenoid derivative--retinal--acts as visual pigment in all animals and as chromophore in bacteriorhodopsin photosynthesis, retinoic acid in animals and abscisic acid in plants serve as hormones. All these functions utilize various properties (mechanical, electronic, stereospecific) of a single structure evolved in bacteria for a single membrane-reinforcing function, thus demonstrating an example of pure evolutionary preadaptation. One of the practical conclusions that can be reached by reviewing uniquely diverse properties and functions of carotenoids is that, when considering possible mechanisms of their effects in organisms (e.g., anticarcinogenic action), all their functional traits should be taken into account.     

Functional Constraints on Insect Immune System Components Govern Their Evolutionary Trajectories

Roles of constraints in shaping evolutionary outcomes are often considered in the contexts of developmental biology and population genetics, in terms of capacities to generate new variants and how selection limits or promotes consequent phenotypic changes. Comparative genomics also recognizes the role of constraints, in terms of shaping evolution of gene and genome architectures, sequence evolutionary rates, and gene gains or losses, as well as on molecular phenotypes. Characterizing patterns of genomic change where putative functions and interactions of system components are relatively well described offers opportunities to explore whether genes with similar roles exhibit similar evolutionary trajectories. Using insect immunity as our test case system, we hypothesize that characterizing gene evolutionary histories can define distinct dynamics associated with different functional roles. We develop metrics that quantify gene evolutionary histories, employ these to characterize evolutionary features of immune gene repertoires, and explore relationships between gene family evolutionary profiles and their roles in immunity to understand how different constraints may relate to distinct dynamics. We identified three main axes of evolutionary trajectories characterized by gene duplication and synteny, maintenance/stability and sequence conservation, and loss and sequence divergence, highlighting similar and contrasting patterns across these axes amongst subsets of immune genes. Our results suggest that where and how genes participate in immune responses limit the range of possible evolutionary scenarios they exhibit. The test case study system of insect immunity highlights the potential of applying comparative genomics approaches to characterize how functional constraints on different components of biological systems govern their evolutionary trajectories.     

The ecosystem and evolutionary contexts of allelopathy

Plants can release chemicals into the environment that suppress the growth and establishment of other plants in their vicinity: a process known as 'allelopathy'. However, chemicals with allelopathic functions have other ecological roles, such as plant defense, nutrient chelation, and regulation of soil biota in ways that affect decomposition and soil fertility. These ecosystem-scale roles of allelopathic chemicals can augment, attenuate or modify their community-scale functions. In this review we explore allelopathy in the context of ecosystem properties, and through its role in exotic invasions consider how evolution might affect the intensity and importance of allelopathic interactions.     

Evolutionary ecology of the interactions between aphids and their parasitoids

Many organisms, including entomopathogenous fungi, predators or parasites, use aphids as ressources. Parasites of aphids are mostly endoparasitoid insects, i.e. insects which lay eggs inside the body of an other insect which will die as a result of their development. In this article, we review the consequences of the numerous pecularities of aphid biology and ecology for their endoparasitoids, notably the Aphidiinae (Hymenoptera: Braconidae). We first examine the various mechanisms used by aphids for defence against these enemies. We then explore the strategies used by aphidiine parasitoids to exploit their aphid hosts. Finally, we consider the responses of both aphids and parasitoids to ecological constraints induced by seasonal cycles and to environmental variations linked to host plants and climate. The fundamental and applied interest of studying these organisms is discussed.     

The evolutionary ecology of insect resistance to plant chemicals

Understanding the diversity of insect responses to chemical pressures (e.g. plant allelochemicals and pesticides) in their local ecological context represents a key challenge in developing durable pest control strategies. To what extent do the resistance mechanisms evolved by insects to deal with the chemical defences of plants differ from those that have evolved to resist insecticides? Here, we review recent advances in our understanding of insect resistance to plant chemicals, with a special emphasis on their underlying molecular basis, evaluate costs associated with each resistance trait, and discuss the ecological and evolutionary significance of these findings.     

Interactions between exotic invasive plants and soil microbes in the rhizosphere suggest that 'everything is not everywhere'

BACKGROUND: The study of soil biota in the context of exotic plant invasions has led to an explosion in our understanding of the ecological roles of many different groups of microbes that function in roots or at the root-soil interface. Part of this progress has been the emergence of two biogeographic patterns involving invasive plants and soil microbes. First, in their non-native ranges invasive plants commonly interact differently with the same soil microbes than native plants. Second, in their native ranges, plants that are invasive elsewhere commonly interact functionally with soil microbes differently in their home ranges than they do in their non-native ranges. These studies pose a challenge to a long-held paradigm about microbial biogeography - the idea that microbes are not limited by dispersal and are thus free from the basic taxonomic, biogeographical and evolutionary framework that characterizes all other life on Earth. As an analogy, the global distribution of animals that function as carnivores does not negate the fascinating evolutionary biogeographic patterns of carnivores. Other challenges to this notion come from new measurements of genetic differences among microbes across geographic boundaries, which also suggest that meaningful biogeographic patterns exist for microorganisms. SCOPE AND CONCLUSIONS: We expand this discussion of whether or not 'everything is everywhere' by using the inherently biogeographic context of plant invasions by reviewing the literature on interactions among invasive plants and the microorganisms in the rhizosphere. We find that these interactions can be delineated at multiple scales: from individual plants to continents. Thus the microbes that regulate major aspects of plant biology do not appear to be exempt from the fundamental evolutionary processes of geographical isolation and natural selection. At the important scales of taxonomy, ecotype and ecosystem functions, the fundamental ecology of invaders and soil microbes indicates that everything might not be everywhere.     

An omics evolutionary perspective on phytophagous insect–host plant interactions in Anastrepha obliqua: a review

Phytophagous insects have a close relationship with their host plants. For this reason, their interactions can lead to important changes in insect population dynamics and evolutionary trajectories. Next generation sequencing (NGS) has provided an opportunity to analyze omics data on a large scale, facilitating the change from a classical genetics approach to a more holistic understanding of the underlying molecular mechanisms of host plant use by insects. Most studies have been carried out on model species in Holarctic and temperate zones. In tropical zones, however, the effects of use of various host plants on evolutionary insect history is less understood. In the current review, we describe how omics methodologies help us to understand phytophagous insect–host plant interactions from an evolutionary perspective, using as example the Neotropical phytophagous insect West Indian fruit fly, Anastrepha obliqua (Macquart) (Diptera: Tephritidae), an economically important fruit crop pest in the Americas. Anastrepha obliqua could adopt a generalist or a specialist lifestyle. We first review the adaptive molecular mechanisms of phytophagous insects to host plants, and then describe the main tools to study phytophagous insect–host plant interactions in the era of omics sciences. The omics approaches will advance the understanding of insect molecular mechanisms and their influence on diversification and evolution. Finally, we discuss the importance of a multidisciplinary approach that integrates the use of omics tools and other, more classical methodologies in evolutionary studies.     

Plant carotenoid cleavage oxygenases and their apocarotenoid products

The oxidative cleavage of carotenoids leads to the production of apocarotenoids and is catalyzed by a family of carotenoid cleavage dioxygenases (CCDs). CCDs often exhibit substrate promiscuity, which probably contributes to the diversity of apocarotenoids found in nature. Biologically and commercially important apocarotenoids include the phytohormone abscisic acid, the visual and signaling molecules retinal and retinoic acid, and the aromatic volatile beta-ionone. Unexpected properties associated with the CCD catalytic products emphasize their role in many aspects of plant growth and development. For instance, CCD7 and CCD8 produce a novel, graft-transmissible hormone that controls axillary shoot growth in plants. Here, CCDs are discussed according to their roles in the biosynthesis of these products. Recent studies regarding their mechanism of action are also addressed.     

雌雄同体植物的性别干扰及其进化意义

雌雄同体植物在传粉和交配过程中两性功能存在潜在的冲突和妥协。除自交和近交衰退外,人们近年来更加强调导致配子浪费和适合度减少的繁殖代价——性别干扰。性别干扰潜在地存在于雌雄同体植物中,尤其是花粉和柱头空间位置接近而又同时成熟的两性花中。该文首先介绍了雌雄同体植物性别干扰的含义及其各种形式,进而用性别干扰理论来解释各种花部性状的适应意义,同时还回顾了植物中关于性别干扰的少数实验证据。该文强调,通常被解释为避免自交的花部机制实际上更大可能是为了避免性别干扰。从更广泛意义上,自交也可以看成是一种形式的性别干扰。