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Host plant quality and fecundity in herbivorous insects.

Host plant quality is a key determinant of the fecundity of herbivorous insects. Components of host plant quality (such as carbon, nitrogen, and defensive metabolites) directly affect potential and achieved herbivore fecundity. The responses of insect herbivores to changes in host plant quality vary within and between feeding guilds. Host plant quality also affects insect reproductive strategies: Egg size and quality, the allocation of resources to eggs, and the choice of oviposition sites may all be influenced by plant quality, as may egg or embryo resorption on poor-quality hosts. Many insect herbivores change the quality of their host plants, affecting both inter- and intraspecific interactions. Higher-trophic level interactions, such as the performance of predators and parasitoids, may also be affected by host plant quality. We conclude that host plant quality affects the fecundity of herbivorous insects at both the individual and the population scale.

Animals↗

Selective toxicity of neonicotinoids attributable to specificity of insect and mammalian nicotinic receptors.

Neonicotinoids, the most important new class of synthetic insecticides of the past three decades, are used to control sucking insects both on plants and on companion animals. Imidacloprid (the principal example), nitenpyram, acetamiprid, thiacloprid, thiamethoxam, and others act as agonists at the insect nicotinic acetylcholine receptor (nAChR). The botanical insecticide nicotine acts at the same target without the neonicotinoid level of effectiveness or safety. Fundamental differences between the nAChRs of insects and mammals confer remarkable selectivity for the neonicotinoids. Whereas ionized nicotine binds at an anionic subsite in the mammalian nAChR, the negatively tipped ("magic" nitro or cyano) neonicotinoids interact with a proposed unique subsite consisting of cationic amino acid residue(s) in the insect nAChR. Knowledge reviewed here of the functional architecture and molecular aspects of the insect and mammalian nAChRs and their neonicotinoid-binding site lays the foundation for continued development and use of this new class of safe and effective insecticides.

Alkaloids↗

The gut bacteria of insects: nonpathogenic interactions.

The diversity of the Insecta is reflected in the large and varied microbial communities inhabiting the gut. Studies, particularly with termites and cockroaches, have focused on the nutritional contributions of gut bacteria in insects living on suboptimal diets. The indigenous gut bacteria, however, also play a role in withstanding the colonization of the gut by non-indigenous species including pathogens. Gut bacterial consortia adapt by the transfer of plasmids and transconjugation between bacterial strains, and some insect species provide ideal conditions for bacterial conjugation, which suggests that the gut is a "hot spot" for gene transfer. Genomic analysis provides new avenues for the study of the gut microbial community and will reveal the molecular foundations of the relationships between the insect and its microbiome. In this review the intestinal bacteria is discussed in the context of developing our understanding of symbiotic relationships, of multitrophic interactions between insects and plant or animal host, and in developing new strategies for controlling insect pests.

Animals↗

Effects of plants genetically modified for insect resistance on nontarget organisms.

Insect resistance, based on Bacillus thuringiensis (Bt) endotoxins, is the second most widely used trait (after herbicide resistance) in commercial genetically modified (GM) crops. Other modifications for insect resistance, such as proteinase inhibitors and lectins, are also being used in many experimental crops. The extensive testing on nontarget plant-feeding insects and beneficial species that has accompanied the long-term and wide-scale use of Bt plants has not detected significant adverse effects. GM plants expressing other insect-resistant proteins that have a broader spectrum of activity have been tested on only a limited number of nontarget species. Little is known about the persistence of transgene-derived proteins in soil, with the exception of Bt endotoxins, which can persist in soil for several months. Bt plants appear to have little impact on soil biota such as earthworms, collembolans, and general soil microflora. Further research is required on the effects of GM plants on soil processes such as decomposition. Assessment of nontarget impacts is an essential part of the risk assessment process for insect-resistant GM plants.

Animals↗

Prostaglandins and other eicosanoids in insects: biological significance.

Prostaglandins and other eicosanoids are oxygenated metabolites of certain polyunsaturated fatty acids. These compounds are well known for their important actions in mammalian physiology and disease. Recent work has revealed the presence and biological actions of eicosanoids in insects and many other invertebrate animals. In insects, eicosanoids mediate cellular immunity to microbial and metazoan challenge. Notably, some infectious organisms secrete factors responsible for impairing host insect immune reactions by inhibiting biosynthesis of eicosanoids. Eicosanoids also act in insect reproductive biology, in ion transport physiology, and in fever response to infection as well as in protein exocytosis in tick salivary glands. Aside from ongoing actions in homeostasis, certain eicosanoid actions occur at crucial points in insect life histories, such as during infectious challenge and important events in reproduction.

Aedes↗

The role of allatostatins in juvenile hormone synthesis in insects and crustaceans.

Allatostatins are pleiotropic neuropeptides for which one function in insects is the inhibition of juvenile hormone synthesis. Juvenile hormone, an important regulator of development and reproduction in insects, is produced by the corpora allata. Mandibular organs, the crustacean homologs of insect corpora allata, produce precursors of juvenile hormone with putatively similar functions. Three types of allatostatins in insects have been isolated: FGLamides, W(X)(6)Wamides, and PISCFs. All act rapidly and reversibly; however, although these types occur in all groups of insects studied, they act as inhibitors of juvenile hormone production in only some groups. Only the FGLamide-type peptides have been isolated in crustaceans, in which they may function to stimulate production of hormone by the mandibular glands, as occurs in early cockroach embryos. Much remains to be learned in order to understand the role of allatostatins in the modulation of hormone production.

Animals↗

Physiology of cold tolerance in insects.

From the available experimental data a relatively clear picture can be established with regard to the physiological importance of some of the mechanisms involved in insect cold hardening. In freeze-avoiding insects, all potent ice-nucleating agents are removed or inactivated, leading to a depression of the supercooling points to about 20 degrees C. Accumulation of polyols causes a further depression with a magnitude of about twice the corresponding melting-point depression. Production of thermal hysteresis factors causes a stabilization of the supercooled state. In freeze-tolerant insects, potent ice-nucleating agents are produced in the extracellular body fluid, ensuring a protective extracellular freezing at a few degrees below zero. Accumulation of polyols causes a steep drop in the lethal temperature, due to a reduction of the amount of ice by a colligative mechanism. However, there is still much to be learned about the mechanisms by which ice-nucleating agents, polyols, and thermal hysteresis agents are acting. Furthermore, the regulatory mechanisms involved in the production and elimination of these components from the body fluid of the insects are not understood. Also, when it comes to the influence of environmental factors, like photoperiod and temperature, there is much to be learned. In addition to giving attention to these topics, future research should be focused on the possible role of other factors in cold hardening such as bound water, dehydration, low-molecular-weight solutes other than polyols, and the biochemical mechanisms forming the basis of the seasonal changes in the cold hardiness of insects.

Adaptation, Physiological↗

The evolution of insect wings and their sensory apparatus.

The development of wings has undoubtedly played a major role in the enormous diversification of insects. New insights into the evolutionary history of insect wings are available from paleontological, physiological and biomechanical studies. A recent hypothesis, derived primarily from paleontological evidence, is that wings arose from leg exites, small flaps associated with proximal leg segments. We present data from studies on physical models that are consistent with this hypothesis. The exites would have been moveable, and measurements on scaled models show that they would have generated aerodynamic lift by unsteady mechanisms associated with vortex shedding. An examination of the sensory structures found on insect wings is also consistent with the interpretation of protowings as leg exites. In addition to mechanosensory bristles, such as are found all over the body, the wings of modern insects carry campaniform sensilla sensitive to cuticular deformation and contact chemoreceptors whose stimulation elicits a feeding response. Both classes of receptors are also found on the legs of modern insects but not on the thorax, favoring the leg exite theory.

Animals↗

Dispersal strategies of phytophagous insects at a local scale: adaptive potential of aphids in an agricultural environment.

BACKGROUND: The spread of agriculture greatly modified the selective pressures exerted by plants on phytophagous insects, by providing these insects with a high-level resource, structured in time and space. The life history, behavioural and physiological traits of some insect species may have evolved in response to these changes, allowing them to crowd on crops and to become agricultural pests. Dispersal, which is one of these traits, is a key concept in evolutionary biology but has been over-simplified in most theoretical studies. We evaluated the impact of the local-scale dispersal strategy of phytophagous insects on their fitness, using an individual-based model to simulate population dynamics and dispersal between leaves and plants, by walking and flying, of the aphid Aphis gossypii, a major agricultural pest, in a melon field. We compared the optimal values for dispersal parameters in the model with the corresponding observed values in experimental trials. RESULTS: We show that the rates of walking and flying disperser production on leaves were the most important traits determining the fitness criteria, whereas dispersal distance and the clustering of flying dispersers on the target plant had no effect. We further show that the effect of dispersal parameters on aphid fitness depended strongly on plant characteristics. CONCLUSION: Parameters defining the dispersal strategies of aphids at a local scale are key components of the fitness of these insects and may thus be essential in the adaptation to agricultural environments that are structured in space and time. Moreover, the fact that the effect of dispersal parameters on aphid fitness depends strongly on plant characteristics suggests that traits defining aphid dispersal strategies may be a cornerstone of host-plant specialization.

Adaptation, Physiological↗

Biosynthesis and secretion of insect lipoprotein: involvement of furin in cleavage of the apoB homolog, apolipophorin-II/I.

The biosynthesis of neutral fat-transporting lipoproteins involves the lipidation of their nonexchangeable apolipoprotein. In contrast to its mammalian homolog apolipoprotein B, however, insect apolipophorin-II/I (apoLp-II/I) is cleaved posttranslationally at a consensus substrate sequence for furin, resulting in the appearance of two apolipoproteins in insect lipoprotein. To characterize the cleavage process, a truncated cDNA encoding the N-terminal 38% of Locusta migratoria apoLp-II/I, including the cleavage site, was expressed in insect Sf9 cells. This resulted in the secretion of correctly processed apoLp-II and truncated apoLp-I. The cleavage could be impaired by the furin inhibitor decanoyl-Arg-Val-Lys-Arg-chloromethyl ketone (decRVKRcmk) as well as by mutagenesis of the consensus substrate sequence for furin, as indicated by the secretion of uncleaved apoLp-II/I-38. Treatment of L. migratoria fat body, the physiological site of lipoprotein biosynthesis, with decRVKRcmk similarly resulted in the secretion of uncleaved apoLp-II/I, which was integrated in lipoprotein particles of buoyant density identical to wild-type high density lipophorin (HDLp). These results show that apoLp-II/I is posttranslationally cleaved by an insect furin and that biosynthesis and secretion of HDLp can occur independent of this processing step. Structure modeling indicates that the cleavage of apoLp-II/I represents a molecular adaptation in homologous apolipoprotein structures. We propose that cleavage enables specific features of insect lipoproteins, such as low density lipoprotein formation, endocytic recycling, and involvement in coagulation.

Amino Acid Chloromethyl Ketones↗

The remarkable conservation of corticotropin-releasing hormone (CRH)-binding protein in the honeybee (Apis mellifera) dates the CRH system to a common ancestor of insects and vertebrates.

CRH-binding protein (CRH-BP) is a key factor in the regulation of CRH signaling; it modulates the bioactivity and bioavailability of CRH and its related peptides. The conservation of CRH-BP throughout vertebrates was only recently demonstrated. Here we report the presence of CRH-BP in the honeybee (Apis mellifera) and other insects. Honeybee CRH-BP resembles previously characterized vertebrate CRH-BP sequences with respect to conserved cysteine residues, gene organization, and overall sequence identity. Phylogenetic analyses confirm the unambiguous orthology of insect and vertebrate CRH-BP sequences. Soon after their discovery, it was noted that insect diuretic hormone-I (DH-I) and its receptor share similarities with the vertebrate CRH family and their receptors. Despite these similarities, demonstration of common ancestry of DH-I and the vertebrate CRH family is still speculative: the mature neuropeptides are short, and their genes differ substantially with regard to the number of coding exons. Moreover, DH and CRH receptors belong to the much larger family of G protein-coupled receptors. In contrast, the unique and conspicuous features of CRH-BP greatly facilitate the establishment of orthology over much larger evolutionary distances. The identification of CRH-BP in insects clearly indicates that this gene predates vertebrates by at least several hundred million years. Moreover, our findings imply that a CRH system is shared by insects and vertebrates alike and, consequently, that it has been present at least since the common ancestor to both phylogenetic lines of proto- and deuterostomians.

Amino Acid Sequence↗

Expression of biologically active human thyrotropin (hTSH) in a baculovirus system: effect of insect cell glycosylation on hTSH activity in vitro and in vivo.

To obtain large amounts of hTSH and to study the role of the N-linked oligosaccharides for its biological activity, hTSH was produced using recombinant baculovirus containing the human alpha-subunit and a hTSH beta-minigene, respectively, both under the control of the polyhedrin promoter. Expression in insect cells was 800-1000 ng/ml, 30-fold higher than in our optimized mammalian transient transfection system using Chinese hamster ovary (CHO) cells (20-50 ng/ml). The in vitro activity of insect-cell expressed hTSH (IC-hTSH) was increased 5-fold compared with CHO-hTSH, judged by the ability to induce cAMP production in CHO cells stably transfected with the hTSH receptor (JP09) and the rat thyroid cell line FRTL-5, as well as growth promotion in FRTL-5 cells. Lectin binding and enzymatic desialylation studies suggested that in contrast to CHO-hTSH, IC-hTSH lacked complex-type oligosaccharides terminating with sialic acid but contained predominantly high mannose-type oligosaccharides. The in vitro activity of CHO-hTSH also increased 5- to 6-fold upon treatment of the hTSH-producing cells with the oligosaccharide processing inhibitors swainsonine and castanospermine, which inhibit formation of complex, terminally sialylated oligosaccharides, and upon enzymatic desialylation. In contrast, insect cell-expression or treatment with processing inhibitors did not affect TSH receptor binding. Despite the higher in vitro activity, IC-hTSH had a much lower in vivo activity than CHO-hTSH, due to rapid clearance from the circulation. In summary, this study shows for the first time that relatively high levels of recombinant hTSH with high in vitro bioactivity can be produced in a baculovirus system. Cell-dependent glycosylation is a major factor that determines the final in vivo biopotency of recombinant glycoproteins, a finding that should be of general relevance for all insect cell-produced glycosylated proteins. Although not suitable for clinical use, highly bioactive recombinant hTSH derived from high expression in insect cells should be useful in defining structure-function relations of hormone analogs.

Animals↗

hunchback is required for suppression of abdominal identity, and for proper germband growth and segmentation in the intermediate germband insect Oncopeltus fasciatus.

Insects such as Drosophila melanogaster undergo a derived form of segmentation termed long germband segmentation. In long germband insects, all of the body regions are specified by the blastoderm stage. Thus, the entire body plan is proportionally represented on the blastoderm. This is in contrast to short and intermediate germband insects where only the most anterior body regions are specified by the blastoderm stage. Posterior segments are specified later in embryogenesis during a period of germband elongation. Although we know much about Drosophila segmentation, we still know very little about how the blastoderm of short and intermediate germband insects is allocated into only the anterior segments, and how the remaining posterior segments are produced. In order to gain insight into this type of embryogenesis, we have investigated the expression and function of the homolog of the Drosophila gap gene hunchback in an intermediate germ insect, the milkweed bug, Oncopeltus fasciatus. We find that Oncopeltus hunchback (Of'hb) is expressed in two phases, first in a gap-like domain in the blastoderm and later in the posterior growth zone during germband elongation. In order to determine the genetic function of Of'hb, we have developed a method of parental RNAi in the milkweed bug. Using this technique, we find that Oncopeltus hunchback has two roles in anterior-posterior axis specification. First, Of'hb is required to suppress abdominal identity in the gnathal and thoracic regions. Subsequently, it is then required for proper germband growth and segmentation. In milkweed bug embryos depleted for hunchback, these two effects result in animals in which a relatively normal head is followed by several segments with abdominal identity. This phenotype is reminiscent to that found in Drosophila hunchback mutants, but in Oncopeltus is generated through the combination of the two separate defects.

Abdomen↗

even-skipped is not a pair-rule gene but has segmental and gap-like functions in Oncopeltus fasciatus, an intermediate germband insect.

The pair-rule gene even-skipped is required for the initiation of metameric pattern in Drosophila. But Drosophila segmentation is evolutionarily derived and is not representative of most insects. Therefore, in order to shed light on the evolution of insect segmentation, homologs of the pair-rule gene even-skipped have been studied in several insect taxa. However, most of these studies have reported the expression eve but not its function. We report the isolation, expression and function of the homolog of Drosophila even-skipped from the intermediate germband insect Oncopeltus fasciatus. We find that in Oncopeltus, even-skipped striped expression initiates in a segmental and not pair-rule pattern. Weak RNAi suppression of Oncopeltus even-skipped shows no apparent pair-rule like phenotype, while stronger RNAi suppression shows deletion of nearly the entire body. These results suggest that in Oncopeltus, even-skipped is not acting as a pair-rule gene. In almost all insects, prior to its striped expression, even-skipped is expressed in a conserved broad gap-like domain but its function has been largely ignored. We find that this early broad domain is required for activation of the gap genes hunchback and Krüppel. Given the large RNAi deletion phenotype and its regulation of hunchback and Krüppel, even-skipped seems to act as an über-gap gene in Oncopeltus, indicating that it may have both upstream and downstream roles in segmentation.

Animals↗

Why is it worth flying at dusk for aquatic insects? Polarotactic water detection is easiest at low solar elevations.

Using 180 degrees field-of-view imaging polarimetry, we measured the reflection-polarization patterns of two artificial surfaces (water-dummies) in the red, green and blue spectral ranges under clear and partly cloudy skies at different solar elevations. The dummies consisted of a horizontal glass pane with a matt black or matt light grey cloth underneath, imitating a dark or bright water body, respectively. Assuming that polarotactic water insects interpret a surface as representing water if the degree of linear polarization of reflected light is higher than a threshold and the deviation of the direction of polarization from the horizontal is lower than a threshold, we calculated the proportion, P, of the artificial surfaces detected polarotactically as water. We found that at sunrise and sunset P is maximal for both water-dummies and their reflection-polarizational characteristics are most similar. From this, we conclude that polarotactic water detection is easiest at low solar elevations, because the risk that a polarotactic insect will be unable to recognize the surface of a dark or bright water body is minimal. This partly explains why many aquatic insect species usually fly en masse at dusk. The daily change in the reflection-polarization pattern of water surfaces is an important visual ecological factor that may contribute to the preference of the twilight period for habitat searching by polarotactic water insects. Air temperature at sunrise is generally low, so dusk is the optimal period for polarotactic aquatic insects to seek new habitats.

Animals↗

The role of discontinuous gas exchange in insects: the chthonic hypothesis does not hold water.

Insects breathe through valved openings (spiracles) in their cuticle. Many insects open and close their spiracles in a cyclic pattern (discontinuous gas-exchange cycles, or DGC). These cycles were observed over half a century ago, their hypothesized function being to minimize loss of water from the tracheal system. However, numerous recent studies have found that respiration accounts for a small fraction of total water loss, and that insects stop performing DGC at times when this pattern would be most useful. Thus, the importance of cyclic gas exchange for water conservation has been challenged. The leading alternative is the chthonic hypothesis, which proposes that DGC originated in insects from hypercapnic (high CO2) environments (e.g. burrows) to aid in release of carbon dioxide. By keeping the spiracles closed, insects would concentrate CO2 and increase the gradient for outward diffusion of CO2. CO2 would be released rapidly when the spiracles opened, and respiratory water loss would be reduced. The chthonic hypothesis therefore predicts that DGC minimizes the ratio of respiratory water loss to CO2 release relative to other modes of gas exchange. We tested the chthonic hypothesis by simultaneously measuring water loss and CO2 release in reproductive females (queens) of the seed-harvester ant Pogonomyrmex barbatus, a burrowing species from North American deserts. Queens used one of three patterns of gas exchange, discontinuous, cyclic and continuous. We resolved the problem of separating cuticular transpiration and respiratory water loss for individuals that used continuous gas exchange by developing a regression method that can be used across all patterns of gas exchange. The ratio of respiratory water loss to CO2 release did not differ among ants using different patterns of gas exchange, in contrast to the expectation of the chthonic hypothesis. Metabolic rate, however, varied with gas-exchange pattern, and was lowest for individuals that used discontinuous gas exchange, intermediate for individuals using cyclic gas exchange, and highest for individuals using continuous gas exchange.

Analysis of Variance↗

Insect navigation en route to the goal: multiple strategies for the use of landmarks

There are at least four distinct ways in which familiar landmarks aid an insect on its trips between nest and foraging site. Recognising scenes: when bees are displaced unexpectedly from their hive to one of several familiar locations, they are able to head in the direction of home as though they had previously linked an appropriate directional vector to a view of the scene at the release site. Biased detours: ants recognise familiar landmarks en route and will correct their path by steering consistently to the left or to the right around them. Aiming at beacons: bees and ants also guide their path by approaching familiar landmarks lying on or close to the direct line between start and finish. Simulations suggest that such mechanisms acting together may suffice to account for the routes taken by desert ants through a landmark-strewn environment: the stereotyped trajectories of individual ants can be modelled by a weighted combination of dead reckoning, biased detours and beacon-aiming. These mechanisms guide an insect sufficiently close to an inconspicuous goal for image matching to be successfully employed to locate it. Insects then move until their current retinal image matches a stored view of the surrounding panorama seen from a vantage point close to the goal. Bees and wasps perform learning flights on their first departure from a site to which they will return. These flights seem to be designed to pick up the information needed for several navigational strategies. Thus, a large portion of the learning flight of a bee leaving a feeder tends to be spent close to the feeder so aiding the acquisition of a view from that vantage point, as is needed for image matching. Bees and social wasps also tend to inspect their surroundings while facing along preferred directions and to adopt similar bearings before landing, thereby making it easy to employ retinotopically stored patterns in image matching. Aiming at beacons, in contrast, requires a landmark to be familiar to the frontal retina. Objects tend to be viewed frontally while the insect circles through arcs centred on the goal. This procedure may help insects to pick out those objects close to the goal that are best suited for guiding later returns.

Journal Article↗

Maintenance of the K+ activity gradient in insect muscle compared in diptera and lepidoptera: contributions of metabolic and exchanger mechanisms

Using a comparative approach, the mechanisms involved in maintenance of the transmembrane K+ activity gradients in the larval body-wall muscles of two insects, Phormia terraenovae (Diptera) and Spodoptera exigua (Lepidoptera), have been investigated. Double-barrelled K+-selective microelectrodes were used to obtain simultaneous measurements of intracellular K+ activity and membrane potential, whilst ordinary microelectrodes were used to monitor input resistance. By application of a variety of general metabolic blockers, the K+ gradients in both P. terraenovae and S. exigua muscle were found to be maintained, at least in part, by a metabolic component. Differences in sensitivity to dinitrophenol of the two insects suggested that the ATP-dependence of maintenance of the K+ gradient was significantly higher in P. terraenovae than in S. exigua. Vanadate sensitivity suggested that both insects possess P-type ATPases. The K+ activity gradient in P. terraenovae muscles was also found to be ouabain-sensitive, indicating the involvement of a Na+/K+-ATPase. In contrast, the K+ gradient in S. exigua muscles proved to be totally insensitive to ouabain but sensitive to amiloride. Application of the H+/K+-ATPase-specific inhibitor SCH 28080 suggested the presence of an H+/K+ pump similar to the mammalian gastric H+/K+-ATPase in the lepidopteran muscles. P. terraenovae muscles, however, were found to be totally insensitive to this inhibitor. Using the anion (Cl-)-dependent transport inhibitors bumetanide and SITS (4-acetamide-4-isothiocyanostilbene-2,2-disulphonic acid), P. terraenovae muscles were shown not to possess a Cl--dependent K+ transport mechanism. In contrast, a bumetanide-sensitive K+/Cl- cotransporter was likely to be involved in maintenance of the K+ gradient in S. exigua muscle. An additional SITS-sensitive Cl-/HCO3- exchanger could also have some indirect involvement in K+ maintenance through regulation of the inward Cl- gradient. The results are integrated in two ionic models, one for each insect, which could account for the bulk of K+ transport in the body-wall muscles of these insects.

Journal Article↗