Yoram Yerushalmi

The physiology of locust phase polymorphism: an update.

The considerable progress made between 1990 and 1997 in locust phase-related research and in understanding the physiology of locust phase polymorphism is reviewed. The traits of locust phases are discussed and it is concluded that there are distinct strain-dependent differences in phase characteristics and their amplitudes even in the same species. Despite some advances, no major break-through was achieved in the putative endocrine control of locust phase polymorphism. Phase-dependent differences in adipokinesis, flight fuels and migration of adult locusts, as well as novel methods in studying aggregation behaviour and activity of hoppers and adults, opened new lines in research of the physiology of locust phase polymorphism. Marked advances were made in phase-related locust pheromone research, revealing, in Schistocerca gregaria, differences between the pheromonal system of the hoppers and that of the adults. These systems turned out to be more complex than previously assumed. Phenylacetonitrile, produced by sexually mature adult males, serving both as an attractant and a mutration-accelerating factor, was identified as the major compound of the adult pheromonal system in S. gregaria. A new aspect of transmission of phase characteristics from parent to progeny through the foam (froth) of the egg pod was revealed. Effects of some plant substances on locust phases were reported. However, no research has yet been published on the aspects of molecular biology of locust phase polymorphism.

Discrete pulses of molting hormone, 20-hydroxyecdysone, during late larval development of Drosophila melanogaster: Correlations with changes in gene activity

Periodic pulses of the insect steroid molting hormone 20‐hydroxyecdysone (20E), acting via its nuclear receptor complex (EcR/USP), control gene expression at many stages throughout Drosophila development. However, during the last larval instar of some lepidopteran insects, subtle changes in titers of ecdysteroids have been documented, including the so‐called “commitment peak.” This small elevation of 20E reprograms the larva for metamorphosis to the pupa. Similar periods of ecdysteroid immunoreactivity have been observed during the last larval instar of Drosophila. However, due to low amplitude and short duration, along with small body size and staging difficulties, their timing and ecdysteroid composition have remained uncertain. Employing a rigorous regimen of Drosophila culture and a salivary gland reporter gene, Sgs3‐GFP, we used RP‐HPLC and differential ecdysteroid RIA analysis to determine whole body titers of 20E during the last larval instar. Three small peaks of 20E were observed at 8, 20, and 28 hr following ecdysis, prior to the well‐characterized large peak around the time of pupariation. The possible regulation of 20E levels by biosynthetic P450 enzymes and the roles of these early peaks in coordinating gene expression and late larval development are discussed. Developmental Dynamics 235:315–326, 2006.

Interaction Between Hormonal Signaling Pathways in Drosophila melanogaster as Revealed by Genetic Interaction Between Methoprene-tolerant and Broad-Complex

Juvenile hormone (JH) regulates insect development by a poorly understood mechanism. Application of JH agonist insecticides to Drosophila melanogaster during the ecdysone-driven onset of metamorphosis results in lethality and specific morphogenetic defects, some of which resemble those in mutants of the ecdysone-regulated Broad-Complex (BR-C). The Methoprene-tolerant (Met) bHLH–PAS gene mediates JH action, and Met mutations protect against the lethality and defects. To explore relationships among these two genes and JH, double mutants were constructed between Met alleles and alleles of each of the BR-C complementation groups: broad (br), reduced bristles on palpus (rbp), and 2Bc. Defects in viability and oogenesis were consistently more severe in rbp Met or br Met double mutants than would be expected if these genes act independently. Additionally, complementation between BR-C mutant alleles often failed when MET was absent. Patterns of BRC protein accumulation during metamorphosis revealed essentially no difference between wild-type and Met-null individuals. JH agonist treatment did not block accumulation of BRC proteins. We propose that MET and BRC interact to control transcription of one or more downstream effector genes, which can be disrupted either by mutations in Met or BR-C or by application of JH/JH agonist, which alters MET interaction with BRC.

Structure–activity relations of the dark-colour-inducing neurohormone of locusts

The dark-colour-inducing effect of several peptides in comparison to that of the dark-colour-inducing neurohormone (DCIN, [His(7)]-corazonin) of locusts was investigated by a bioassay based on nymphs of a DCIN-deficient albino mutant of Locusta migratoria. The study was aimed at elucidating the active part of the DCIN and to explore the contribution of its amino acids to the activity. Graded doses of all peptides were injected in oil. [Arg(7)]-corazonin and DCIN were equally effective. Certain arthropod neuropeptides having the -SXGW- partial sequence (a part of the DCIN and of [Arg(7)]-corazonin; X=His and X=Arg, respectively) yielded the following findings: Scg-AKH-II (adipokinetic hormone II of Schistocerca gregaria X=Thr), Grb-AKH ( adipokinetic hormone of Gryllus bimaculatus X=Thr) and RPCH (red pigment concentrating hormone of crustaceans X=Pro) evoked a moderate darkening response, but Lom-AKH-II (adipokinetic hormone II of L. migratoria X=Ala) was ineffective. Step by step shortening of the sequence of the DCIN at the N-terminal, from pGlu-3-11DCIN to pGlu-9-11DCIN, resulted in a decreasing activity, but even pGlu-9-11DCIN induced a weak response with high doses. Shortening of the DCIN from the C-terminal revealed a moderate activity of 1-7DCIN-NH(2) and a weak activity of 1-5DCIN-NH(2). An octadecapeptide which induces dark colour in moth larvae, having the pentamer FTPRL-NH(2) at its C-terminal, evoked no darkening in the albino locusts. We conclude that although the -SXGW- partial sequence has some role in induction of darkening, for obtaining maximal effect the whole sequence of the DCIN (or of [Arg(7)]-corazonin) is necessary.

Emerging model systems in evo-devo: horned beetles and the origins of diversity

SUMMARY Horned beetles and beetle horns are emerging as a model system suited to address fundamental questions in evolutionary developmental biology. Here we briefly review the biology of horned beetles and highlight the unusual opportunities they provide for evo‐devo research. We then summarize recent advances in the development of new approaches and techniques that are now available to scientists interested in working with these organisms. We end by discussing ways to implement and combine these new approaches to explore new frontiers in evo‐devo research previously unavailable to reseachers working outside traditional model organisms.

Phase polymorphism in Locusta migratoria: the relative effects of geographical strains and albinism on morphometrics

An albino strain, which had originated from Okinawa, Japan, and a normally coloured strain, which had originated from West Africa, have been used to study density‐dependent morphometric phase characteristics and their changes in adults of the migratory locust, Locusta migratoria (L.). By repeated crossings we also obtained congenic albinos and normal phenotypes and investigated their morphometrics with increasing West African genome, eventually reaching 99.6% West African and 0.4% Okinawa gene pool. The data were analysed by the classical morphometric ratios (F/C and E/F; F = length of the hind femur, C = maximum width of the head, E = length of the fore wings), as well as by canonical discriminant (multivariate) analysis. The latter was based on measurements of F, C and E (as above), as well as of M (minimum width of the pronotum) and H (maximum height of the pronontum). Okinawa albinos showed more solitarious morphometrics and a smaller amplitude of morphometric phase change than West African normal phenotypes. Both the morphometric ratios and the canonical discriminant analysis demonstrated clearly that these differences were caused primarily by the strain (Okinawa vs. West African). However, the pigmentation (albino vs. normal colouration) also affected morphometric phase differences; albinos showed more solitarious morphometrics and somewhat more restricted morphometric phase change than congenic normal phenotypes. The effect of the pigmentation was considerably smaller than that of the strain. The results refute Noltes claim that albino locusts constitute an extreme solitarious phase, even under crowding. However, Noltes less extreme claim, that albino locusts have more solitarious morphometrics than normally coloured locusts, is validated by the present results.

The response of a homochrome grasshopper, Oedipoda miniata, to the dark-colour-inducing neurohormone (DCIN) of locusts

The effect of the dark-colour-inducing neurohormone (DCIN=[His(7)]-corazonin) of locusts was investigated in nymphs of the grasshopper, Oedipoda miniata, which exhibit strong homochromy, but neither green-brown, nor phase colour polymorphism. Graded doses of synthetic DCIN were injected in 1 µl of olive oil into 0-24-h-old penultimate nymphs. DCIN induced dose-dependent darkening in the recipients, seen 4 days after injection (still in the penultimate nymphal instar), as well as in the subsequent last-instar nymphs. The dose-range obtained between discernible and maximum (almost black) darkening extended over three orders of magnitude, from 1 pmol to 1 nmol of DCIN. In spite of the darkening observed already in the penultimate nymphal instar, the exuviae in the moult from this to the last nymphal instar did not show marked dark patches. However, the exuviae of the next moult (from last-instar nymph to adult) showed such dark patches, which increased with the increase of the dose, revealing the presence of exocuticular melanin. We conclude that DCIN, or a very close peptide, is the endogenous hormone which plays the major or sole role in the control of homochromy of O. miniata and possibly also of other acridids which exhibit homochromy.

The dark-colour-inducing neurohormone of locusts in relation to an albino mutant of Schistocerca gregaria

In the albino mutant of an Okinawa strain of Locusta migratoria (L.) (Orthoptera: Acrididae), albinism is caused by the absence of the dark‐colour‐inducing neurohormone (DCIN), which is present in the corpora cardiaca (CC) of normally coloured phenotypes. This study tests whether the absence of DCIN is responsible for albinism in an albino mutant of another locust, Schistocerca gregaria (Forsk.) (Orthoptera: Acrididae). This seemed feasible because a single Mendelian unit controls albinism in both species. However, implantation of CC, or injection of an extract of CC, from albino donors of S. gregaria, induce dark coloration in crowded nymph recipients of the Okinawa albino mutant of L. migratoria, as effectively as do implanted CC, or injections of extract of CC, from normal phenotype donors of S. gregaria. Therefore, DCIN is present in the albino mutant of S. gregaria, and consequently, the albinism in this mutant is not caused by its absence.

Locust research in the age of model organisms: introduction to the special issue in honor of M.P. Pener's 80th birthday.

This special issue of the Journal of Insect Physiology is devoted to locust research in the age of model organisms. Prospective authors with diverse research approaches were invited to contribute and we are delighted by the enthusiastic response. All submissions, including broad reviews, targeted reviews, and experimental papers, were peer-reviewed and, in most cases, revised prior to publication. The result is a superb collection of articles, representing a wide variety of research areas with one thing in common: the use of locusts, either as the major research animal or as a model. The goal of collecting these works was to provide a venue through which a panorama of cutting-edge locust research could emerge, demonstrating that even nowadays, when research tends to focus increasingly on a few selected so-called model organisms, the locust has not lost its appeal as an outstanding preparation for physiological studies in various fields, from endocrinology to neurobiology, integrative physiology and biochemistry. Following is a short review of the 21 contributions included in this special issue, divided into five themes or general subjects. Locomotion and neural control: The locust’s superb capacity for flight has always attracted much research (Fig. 1); covering behavioral studies as well as various aspects of neural and neuroendocrine control. The work by Wilson (1961), focusing on the neural mechanisms for the generation of locust flight, has served as a landmark in our understanding of rhythmic behaviors and the central pattern-generating circuits that control them. In their review, Ayali and Lange present the locust as a unique preparation in which central pattern generator networks have been suggested and studied in practically all aspects of the animal’s behavior. These include flight as well as other means of locomotion, respiration, feeding and molting, oviposition and more. Extensive laboratory studies have been carried out on flight fuels, their utilization and mobilization in locusts. Van der Horst and Rodenburg focus on hormonal regulation of lipid mobilization and transport during locust flight activity. This updated review presents the locust as a broad and profound research model for integrative physiology and biochemistry, while also acknowledging its limitations as such. Much work related to neurohormones and flight has been focused on the role of the biogenic amine octopamine. Verlinden et al. provide an excellent and timely reminder of the various roles of octopamine in multiple physiological and behavioral processes in locusts and other arthropods. Octopamine exerts its effects by binding to specific receptor proteins that belong to the superfamily

Age‐dependent response of adults of a homochrome grasshopper, Oedipoda miniata, to the dark‐colour‐inducing neurohormone (DCIN) of locusts

Abstract The effect of the dark‐colour‐inducing neurohormone (DCIN = [His7]‐corazonin) of locusts was investigated in field‐collected young and old adults of the grasshopper, Oedipoda miniata (Pallas) (Orthoptera: Acrididae). This species shows homochromy, but neither green‐brown, nor phase‐dependent colour polymorphism. By injecting graded doses of synthetic DCIN in 2 µL of olive oil, young adults were tested within a week of their last moult, and old adults 3.5 months later, a few weeks before natural termination of their reproductive summer diapause. Colour changes were followed for 28 days after injection. Darkening of the young adults was considerable, but their response to DCIN is more moderate than that of conspecific nymphs, by exhibiting a higher threshold, slower response and weaker maximal response. Old adults also show a clear effect, but their response is even slower and less marked than the response of the young adults. It is concluded that the response to DCIN decreases from nymphs to young adults and it is further decreasing with ageing of the adults.