Behavioural manipulation in a grasshopper harbouring hairworm: a proteomics approach

The parasitic Nematomorph hairworm, Spinochordodes tellinii (Camerano) develops inside the terrestrial grasshopper, Meconema thalassinum (De Geer) (Orthoptera: Tettigoniidae), changing the insect's responses to water. The resulting aberrant behaviour makes infected insects more likely to jump into an aquatic environment where the adult parasite reproduces. We used proteomics tools (i.e. two-dimensional gel electrophoresis (2-DE), computer assisted comparative analysis of host and parasite protein spots and MALDI-TOF mass spectrometry) to identify these proteins and to explore the mechanisms underlying this subtle behavioural modification. We characterized simultaneously the host (brain) and the parasite proteomes at three stages of the manipulative process, i.e. before, during and after manipulation. For the host, there was a differential proteomic expression in relation to different effects such as the circadian cycle, the parasitic status, the manipulative period itself, and worm emergence. For the parasite, a differential proteomics expression allowed characterization of the parasitic and the free-living stages, the manipulative period and the emergence of the worm from the host. The findings suggest that the adult worm alters the normal functions of the grasshopper's central nervous system (CNS) by producing certain 'effective' molecules. In addition, in the brain of manipulated insects, there was found to be a differential expression of proteins specifically linked to neurotransmitter activities. The evidence obtained also suggested that the parasite produces molecules from the family Wnt acting directly on the development of the CNS. These proteins show important similarities with those known in other insects, suggesting a case of molecular mimicry. Finally, we found many proteins in the host's CNS as well as in the parasite for which the function(s) are still unknown in the published literature (www) protein databases. These results support the hypothesis that host behavioural changes are mediated by a mix of direct and indirect chemical manipulation.     

Entomopathogenic fungi and insect behaviour: from unsuspecting hosts to targeted vectors

The behavioural response of an insect to a fungal pathogen will have a direct effect on the efficacy of the fungus as a biological control agent. In this paper we describe two processes that have a significant effect on the interactions between insects and entomopathogenic fungi: (a) the ability of target insects to detect and avoid fungal pathogens and (b) the transmission of fungal pathogens between host insects. The behavioural interactions between insects and entomopathogenic fungi are described for a variety of fungal pathogens ranging from commercially available bio-pesticides to non-formulated naturally occurring pathogens. The artificial manipulation of insect behaviour using dissemination devices to contaminate insects with entomopathogenic fungi is then described. The implications of insect behaviour on the use of fungal pathogens as biological control agents are discussed.     

The smallest insects evolve anucleate neurons

The smallest insects are comparable in size to unicellular organisms. Thus, their size affects their structure not only at the organ level, but also at the cellular level. Here we report the first finding of animals with an almost entirely anucleate nervous system. Adults of the smallest flying insects of the parasitic wasp genus Megaphragma (Hymenoptera: Trichogrammatidae) have only 339-372 nuclei in the central nervous system, i.e., their ganglia, including the brain, consist almost exclusively of processes of neurons. In contrast, their pupae have ganglia more typical of other insects, with about 7400 nuclei in the central nervous system. During the final phases of pupal development, most neuronal cell bodies lyse. As adults, these insects have many fewer nucleated neurons, a small number of cell bodies in different stages of lysis, and about 7000 anucleate cells. Although most neurons lack nuclei, these insects exhibit many important behaviors, including flight and searching for hosts.     

Vision in flying insects

Vision guides flight behaviour in numerous insects. Despite their small brain, insects easily outperform current man-made autonomous vehicles in many respects. Examples are the virtuosic chasing manoeuvres male flies perform as part of their mating behaviour and the ability of bees to assess, on the basis of visual motion cues, the distance travelled in a novel environment. Analyses at both the behavioural and neuronal levels are beginning to unveil reasons for such extraordinary capabilities of insects. One recipe for their success is the adaptation of visual information processing to the specific requirements of the behavioural tasks and to the specific spatiotemporal properties of the natural input.     

Peculiarities of the brain organization and fine structure in small insects related to miniaturization. 2. The smallest Hymenoptera (Mymaridae, Trichogrammatidae)

The organization and fine structure of the brain in the smallest parasitic wasps, Anaphes flavipes and Trichogramma evanescens, were studied using serial histological sections as well as TEM and computer-assisted 3D reconstructions. The data on the number and size of neurons in the brain of Mymaridae and Trichogrammatidae were obtained. They confirm and supplement the hypothesis about the factors limiting miniaturization of insects. The organization of the brain of the smallest parasitic wasps is compared with that of Hemiptarsenus sp. (Eulophidae), a large representative of Chalcidoidea. Some brain areas reveal strong allometry. The relative volume of the antennal lobes, lateral protocerebrum, and protocerebral bridge increases significantly as the body size decreases. Miniaturization is accompanied by an increase in the relative brain volume and changes in the spatial orientation of some brain structures. The number and size of neurons of A. flavipes and T. evanescens are significantly different from those of large representatives of Chalcidoidea, but similar to those of other tiny insects. Miniaturization of the nervous system is limited by the size of neurons and the diameter of axons.     

Behavioural syndromes and social insects: personality at multiple levels

Animal personalities or behavioural syndromes are consistent and/or correlated behaviours across two or more situations within a population. Social insect biologists have measured consistent individual variation in behaviour within and across colonies for decades. The goal of this review is to illustrate the ways in which both the study of social insects and of behavioural syndromes has overlapped, and to highlight ways in which both fields can move forward through the synergy of knowledge from each. Here we, (i) review work to date on behavioural syndromes (though not always referred to as such) in social insects, and discuss mechanisms and fitness effects of maintaining individual behavioural variation within and between colonies; (ii) summarise approaches and principles from studies of behavioural syndromes, such as trade‐offs, feedback, and statistical methods developed specifically to study behavioural consistencies and correlations, and discuss how they might be applied specifically to the study of social insects; (iii) discuss how the study of social insects can enhance our understanding of behavioural syndromes—research in behavioural syndromes is beginning to explore the role of sociality in maintaining or developing behavioural types, and work on social insects can provide new insights in this area; and (iv) suggest future directions for study, with an emphasis on examining behavioural types at multiple levels of organisation (genes, individuals, colonies, or groups of individuals).     

Learning in honeybees: from molecules to behaviour

Studies in a variety of organisms as diverse as molluscs, insects, birds and mammals have shown that memories can exist in a variety of temporal domains ranging from short-term memories in the range of minutes to long-term memories lasting a lifetime. While transient covalent modifications of proteins underlie short-term memory, the formation of long-term memory requires gene expression and protein synthesis. Different intracellular signalling cascades have been implicated in distinct aspects of learning and memory formation. Little is known however, about how learning in intact animals is related to the modulation of these signalling cascades and how this contributes to distinct neuronal and behavioural changes in vivo. Associative learning in the honeybee provides the opportunity to study processes of memory formation by analysing its progression through different phases, across levels of behaviour, neural circuits, and cellular signalling pathways. The findings reveal evidence that various cellular signalling pathways in the neuronal circuit of distinct brain areas play a role in different processes during learning and memory formation.     

Early steps in building the insect brain: neuroblast formation and segmental patterning in the developing brain of different insect species

In insects, morphological, molecular and genetic studies have provided a detailed insight into the ontogenetic processes that shape the ventral nerve cord. On the other hand, owing to its complexity and less obvious segmental composition, the knowledge about the development of the brain is still fragmentary. A promising approach towards gaining insight into fundamental processes underlying brain development is the comparison of embryonic brain development among different insect species. However, so far such comparative analyses are scarce. In this review, we summarize and compare data on the early steps in brain formation in different hemi- and holometabolous insects. We show that basic aspects of the spatial and temporal development of the embryonic brain neuroblast pattern are conserved among insects. Furthermore, we compare the number and proliferation patterns of neuroblasts related to major neuropil structures such as mushroom bodies, central complex, and antennal lobe. Finally, comparing the expression patterns of engrailed in different species, and considering new data from Drosophila melanogaster, we discuss the segmental organization of the insect brain.     

昆虫记忆的形成机制及生态适应性

介绍近十几年来利用蜜蜂Apis mellifera L.、果蝇Drosophila melanogaster Meigen和寄生蜂等昆虫对学习和记忆行为研究的成果。这些研究表明昆虫的记忆形成是多阶段的。从短时记忆向长时记忆的形成过程中,cAMP反应元件结合蛋白(cAMP response element-bindingprotein,CREB)起重要的作用;而蘑菇体是学习、记忆形成的主要位点。已有的研究还表明昆虫记忆的动态是适应于不同物种的生态学需要。这些研究为探索昆虫和其它动物记忆的形成和生态适应性提供了理论基础。     

Types of parasitism by larvae of water mites (Acari: Hydrachnellae)

The larvae of Hydrachnellae can be divided into three types, based on the habits of the parasitic (larval) stage in the life cycle. (1) The larvae do not leave the water. Parasitizing aquatic insects, they are permanently submerged and wet. Only some species of the family Hydrachnidae belong to this type. (2) The larvae leave the water. The parasitic phase occurs either on insects living on the water surface, or in the air stores of aquatic insects. In both cases the larvae are not in direct contact with water during the parasitic phase. The Limnocharidae, Eylaidae and some species of the Hydrachnidae belong to this type. (3) The larvae parasitize insects which live in the air and which can leave the direct proximity to water; therefore it may be difficult for the mite larva to return to water. Except for the Hydrachnidae, Limnocharidae and Eylaidae, all families of the Hydrachnellae belong to this type.     

Bumble-bee foragers infected by a gut parasite have an impaired ability to utilize floral information

Parasitic infection can influence a variety of behavioural mechanisms in animals, but little is known about the effects of infection on the cognitive processes underlying ecologically relevant behaviours. Here, we examined whether parasitic infection alters cognitive aspects of foraging in a social insect, the bumble-bee (Bombus impatiens). In controlled experiments, we assessed the ability of foraging bees to discriminate rewarding from non-rewarding flowers on the basis of colour and odour. We found that natural and experimental infection by a protozoan parasite (Crithidia bombi, which lives exclusively within the gut tract), impaired the ability of foragers to learn the colour of rewarding flowers. Parasitic infection can thus disrupt central nervous system pathways that mediate cognitive processes in bumble-bees and as a consequence, can reduce their ability to monitor floral resources and make economic foraging decisions. It is postulated that this infection-induced change to cognitive function in bumble-bees is the result of communication between immune and nervous systems. Parasitized animals, including invertebrates, can therefore show subtle behavioural changes that are nonetheless ecologically significant and reflect complex mechanisms.     

Plant volatiles induced by herbivore egg deposition affect insects of different trophic levels

Plants release volatiles induced by herbivore feeding that may affect the diversity and composition of plant-associated arthropod communities. However, the specificity and role of plant volatiles induced during the early phase of attack, i.e. egg deposition by herbivorous insects, and their consequences on insects of different trophic levels remain poorly explored. In olfactometer and wind tunnel set-ups, we investigated behavioural responses of a specialist cabbage butterfly (Pieris brassicae) and two of its parasitic wasps (Trichogramma brassicae and Cotesia glomerata) to volatiles of a wild crucifer (Brassica nigra) induced by oviposition of the specialist butterfly and an additional generalist moth (Mamestra brassicae). Gravid butterflies were repelled by volatiles from plants induced by cabbage white butterfly eggs, probably as a means of avoiding competition, whereas both parasitic wasp species were attracted. In contrast, volatiles from plants induced by eggs of the generalist moth did neither repel nor attract any of the tested community members. Analysis of the plant's volatile metabolomic profile by gas chromatography-mass spectrometry and the structure of the plant-egg interface by scanning electron microscopy confirmed that the plant responds differently to egg deposition by the two lepidopteran species. Our findings imply that prior to actual feeding damage, egg deposition can induce specific plant responses that significantly influence various members of higher trophic levels.     

Age and social experience induced plasticity across brain regions of the paper wasp Polistes fuscatus

Developmental studies of brain volumes can reveal which portions of neural circuits are sensitive to environmental inputs. In social insects, differences in relative investment across brain regions emerge as behavioural repertoires change during ontogeny or as a result of experience. Here, we test the effects of maturation and social experience on morphological brain development in Polistes fuscatus paper wasps, focusing on brain regions involved in visual and olfactory processing. We find that mature wasps regardless of social experience have relatively larger brains than newly emerged wasps and this difference is driven by changes to mushroom body calyx and visual regions but not olfactory processing neuropils. Notably, social wasps invest more in the anterior optic tubercle (AOT), a visual glomerulus involved in colour and object processing in other taxa, relative to other visual integration centres the mushroom body calyces compared with aged socially naive wasps. Differences in developmental plasticity between visual and olfactory neuropil volumes are discussed in light of behavioural maturation in paper wasps, especially as it relates to social recognition. Previous research has shown that P. fuscatus need social experience to develop specialized visual processing of faces, which is used to individually recognize conspecifics. The present study suggests that the AOT is a candidate brain region that could mediate facial processing in this species.     

Food behaviour and obesity: insights from decision neuroscience

Neuroimaging allows to estimate brain activity when individuals are doing something. The location and intensity of this estimated activity provides information on the dynamics and processes that guide choice behaviour and associated actions that should be considered a complement to behavioural studies. Decision neuroscience therefore sheds new light on whether the brain evaluates and compares alternatives when decisions are made, or if other processes are at stake. This work helped to demonstrate that the situations faced by individuals (risky, uncertain, delayed in time) do not all have the same (behavioural) complexity, and are not underlined by activity in the cerebral networks. Taking into account brain dynamics of people (suffering from obesity or not) when making food consumption decisions might allow for improved strategies in public health prevention, far from the rational choice theory promoted by neoclassical economics.     

Changes in base composition bias of nuclear and mitochondrial genes in lice (Insecta: Psocodea)

While it is well known that changes in the general processes of molecular evolution have occurred on a variety of timescales, the mechanisms underlying these changes are less well understood. Parasitic lice (“Phthiraptera”) and their close relatives (infraorder Nanopsocetae of the insect order Psocodea) are a group of insects well known for their unusual features of molecular evolution. We examined changes in base composition across parasitic lice and bark lice. We identified substantial differences in percent GC content between the clade comprising parasitic lice plus closely related bark lice (=Nanopsocetae) versus all other bark lice. These changes occurred for both nuclear and mitochondrial protein coding and ribosomal RNA genes, often in the same direction. To evaluate whether correlations in base composition change also occurred within lineages, we used phylogenetically controlled comparisons, and in this case few significant correlations were identified. Examining more constrained sites (first/second codon positions and rRNA) revealed that, in comparison to the other bark lice, the GC content of parasitic lice and close relatives tended towards 50 % either up from less than 50 % GC or down from greater than 50 % GC. In contrast, less constrained sites (third codon positions) in both nuclear and mitochondrial genes showed less of a consistent change of base composition in parasitic lice and very close relatives. We conclude that relaxed selection on this group of insects is a potential explanation of the change in base composition for both mitochondrial and nuclear genes, which could lead to nucleotide frequencies closer to random expectation (i.e., 50 % GC) in the absence of any mutation bias. Evidence suggests this relaxed selection arose once in the non-parasitic common ancestor of Phthiraptera + Nanopsocetae and is not directly related to the evolution of the parasitism in lice.     

云南昆明地区糠片盾蚧的天敌昆虫种类初报

于 2 0 0 1～ 2 0 0 2年在昆明和曲靖对危害桃、苹果和梨树上糠片盾蚧的天敌昆虫种类进行了调查。结果显示 ,糠片盾蚧寄生性天敌昆虫有 7种 ,隶属于膜翅目小蜂总科的跳小蜂科和蚜小蜂科 ,其中 ,桑盾蚧黄蚜小蜂Aphytisproclia (Walker)是糠片盾蚧的主要寄生蜂 ,其数量占寄生蜂总数的 79 7%;捕食性天敌昆虫有 3种 ,隶属于鞘翅目、瓢虫科 ,其中 ,二双斑唇瓢虫ChilocorusbijugusMulsant为糠片盾蚧的主要捕食性天敌昆虫     

Is the composition of phytophagous insects and parasitic fungi among trees predictable?

We explore the relationship between the pairwise similarity of assemblages of exploiters (phytophagous insects and parasitic fungi) and pairwise genetic distance, range overlap, niche overlap as well as habitat overlap of host trees. Presence of exploiters was extracted from published literature for 23 tree genera occurring in central Europe (6164 host records of phytophagous insects and 860 host records of parasitic fungi). Across all pairs of tree genera, we found a strong negative correlation between the pairwise similarity of assemblages and genetic distances of hosts. This close correlation is due to deep differences in the composition of assemblages on coniferous and deciduous tree genera. Range, niche and habitat overlap were always of much less importance than genetic distance to explain the variation of pairwise similarity of assemblages of exploiters, although some correlations were significant. Therefore in general host switches of exploiters between related hosts are more important that host switches between hosts co-occurring in the same habitat. We found a robust relationship of the pairwise similarity of assemblages of insects and the pairwise similarity of assemblages of fungi which points to the possibility that insects are vectors for parasitic fungi which promotes correlated switches of insects and fungi.     

龙眼园节肢动物群落结构及其时空格局

通过对龙眼园节肢动物群落结构及时空格局12次的调查,结果表明,龙眼园节肢动物群落由2纲21目127科334种组成,其中昆虫纲的植食性昆虫8目69科159种（类群）;腐生性昆虫4目9科9种（类群）;捕食性昆虫7目20科47种（类群）;寄生性昆虫1目9科11种（类群）;蛛形纲1目20科108种。各亚群落个体数分别为：植食性昆虫占31．21％,腐生性昆虫占27．15％,捕食性昆虫占25．49％,寄生性昆虫占1．28％,蜘蛛占14．87％;种类数则分别占47．60％,2．69％,14．07％,3．29％,32．34％。群落结构分层明显,树冠上下层皆有的优势类群是黄立毛蚁（Paratrechina flavipes）,多色金蝉蛛（Phintella vesicolor）、啮虫科（Psocidae）,小蜂总科（Chalcidoidea）。树冠上层的优势类群有白蛾蜡蝉（Lawana imitata）,荔蝽（Tessaratoma papillosa）,龙眼角颊木虱（Cornegenapsylla sinica）,折翅蠊科（Blaberidae）,姬蜂科（Ichneumonidae）,下层有矢尖蚧（Unaspis citri）,黑刺粉虱（Aleurocanthus spiniferus）,蚊总科（Culicoidea）,长角跳虫科（Entomobryidae）,细蜂总科（Proctotrupoidea）,咸丰球蛛（Theridion xianfengensis）和灵川丽蛛（Chrysso lingchuanensis）和常见类群小叶蝉（Japananus spp．）。地面杂草层优势类群是比罗举腹蚁（Crematogaster biroi）,黄斑弓背蚁（Camponotus albosparsus）,哀弓背蚁（Camponotus dolendus）,斜纹猫蛛（Oxyopes sertatus）,长角跳虫科,蚊总科,小蜂总科,茧蜂科（Braconidae）,姬蜂科和常见类群幼豹蛛（Pardosa pusiola）。树冠上、下层的植食性和寄生性昆虫亚群落的相似性小,捕食性昆虫和蜘蛛的相似性大。龙眼树冠上的5个亚群落的数量时间格局从1月至10月份是由低到高,然后逐渐下降,其中蜘蛛亚群落的数量波动较平缓,高峰在4月份,其余的群落数量季节波动较大,植食性昆虫数量最高峰在5月份,捕食性昆虫的数量高峰在8月份,寄生性昆虫数量高峰在5月份和7月份。地面杂草层的群落数量波动比树冠层平缓。植食性昆虫与蜘蛛的数量相关显著。     

Pyrokinin neuropeptides in a crustacean. Isolation and identification in the white shrimp Penaeus vannamei.

Identification of substances able to elicit physiological or behavioural processes that are related to reproduction would greatly contribute to the domestication of commercially important crustaceans that do not reproduce easily in captivity. Crustaceans are thought to release urine signals used for chemical communication involved in courtship behaviour. In contrast to insects, very little is known about the endocrinological processes underlying this phenomenon. Therefore, an extract of 3500 central nervous systems of female white shrimp Penaeus vannamei was screened for myotropic activity in order to purify pyrokinin-like peptides that belong to the pyrokinin/PBAN neuropeptide family. Members of this family regulate reproductive processes in insects, including pheromone biosynthesis. Purification of these pyrokinins was achieved by a combination of reversed-phase and normal-phase chromatography. Subsequent characterization by mass spectrometry, Edman degradation and peptide synthesis resulted in the elucidation of two novel peptides. Pev-PK 1 has the primary sequence DFAFSPRL-NH(2) and a second peptide (Pev-PK 2) is characterized as the nonapeptide ADFAFNPRL-NH(2). Pev-PK 1 contains the typical FXPRL-NH(2) (X = G, S, T or V) C-terminal sequence that characterizes members of the versatile pyrokinin/PBAN family. Pev-PK 2 displays an Asn residue at the variable X position of the core pyrokinin sequence. These crustacean pyrokinins are the first to be found in a noninsect. The synthetic peptides display myotropic activity on the Leucophaea maderae as well as on the Astacus leptodactylus hindgut.     

Priming by airborne signals boosts direct and indirect resistance in maize

Plants counteract attack by herbivorous insects using a variety of inducible defence mechanisms. The production of toxic proteins and metabolites that instantly affect the herbivore's development are examples of direct induced defence. In addition, plants may release mixtures of volatile organic compounds (VOCs) that indirectly protect the plant by attracting natural enemies of the herbivore. Recent studies suggest that these VOCs can also prime nearby plants for enhanced induction of defence upon future insect attack. However, evidence that this defence priming causes reduced vulnerability to insects is sparse. Here we present molecular, chemical and behavioural evidence that VOC-induced priming leads to improved direct and indirect resistance in maize. A differential hybridization screen for inducible genes upon attack by Spodoptera littoralis caterpillars identified 10 defence-related genes that are responsive to wounding, jasmonic acid (JA), or caterpillar regurgitant. Exposure to VOCs from caterpillar-infested plants did not activate these genes directly, but primed a subset of them for earlier and/or stronger induction upon subsequent defence elicitation. This priming for defence-related gene expression correlated with reduced caterpillar feeding and development. Furthermore, exposure to caterpillar-induced VOCs primed for enhanced emissions of aromatic and terpenoid compounds. At the peak of this VOC emission, primed plants were significantly more attractive to parasitic Cotesia marginiventris waSPS. This study shows that VOC-induced priming targets a specific subset of JA-inducible genes, and links these responses at the molecular level to enhanced levels of direct and indirect resistance against insect attack.