Effect of [His7]-corazonin on the number of antennal sensilla in Locusta migratoria

Abstract.  Certain types of antennal sensilla are known to be more abundant in solitarious individuals than in gregarious ones in the migratory locust, Locusta migratoria . We tested the hypothesis that injection of a neurohormone, [His⁷]-corazonin, into isolated-reared nymphs of this species mimics the effect of crowding on the frequencies of various types of antennal sensilla. One nmol of the hormone was injected into nymphs on two occasions, on the third days of the second and third stadia, respectively. Upon adult emergence, the numbers of different types of sensilla on the eighth antennal segment were compared with those of oil-injected controls. [His⁷]-corazonin did not influence the numbers of basiconic sensilla type A, basiconic sensilla type B and trichoid sensilla significantly compared to oil-injected controls. However, the number of coeloconic sensilla was reduced significantly by the hormone injections. Because the length of the antennal segment was not affected by the hormone injection, it appears that the hormone influenced the development of coeloconic sensilla. The results support the hypothesis tested and are consistent with the idea that [His⁷]-corazonin plays an important role in the control of phase polymorphism in L. migratoria .     

Hormonal control of body-color polymorphism in Locusta migratoria: interaction between

The effects of injection of [His(7)]-corazonin and juvenile hormone (JH) III on the body color in L. migratoria were investigated using albino and normal (pigmented) nymphs. Most albino nymphs turned green in the fourth instar if injected with JH III during the last 2 days of the previous instar. When albino third instar nymphs injected with 10 pmol of [His(7)]-corazonin on different days were treated with 100 μg of JH III on day 3.5, they developed various body colors in the following nymphal instar: those injected with [His(7)]-corazonin during the first 2 days developed very dark green or black color, whereas some of those injected after this period turned green and their legs and ventral side of the body were variously pigmented, the coloration being similar to green solitary individuals often found in the field. Field-collected brown solitary nymphs injected with 1 nmol of [His(7)]-corazonin and kept individually, turned reddish without any black spots in the following nymphal instar when the ecdysis occurred within 1 day after injection. Injection of [His(7)]-corazonin 2 days before the following ecdysis induced black patterns on an orange background color, the coloration characteristic of gregarious forms. Similar injections into field-collected green solitary nymphs also induced black patterns but the rest of their body remained green. These results may indicate that the temporal changes in the hemolymph titers of [His(7)]-corazonin and JH play an important role in the control of body-color polymorphism in this locust.     

The role of

The effect of [His(7)]-corazonin on the body color in Locusta migratoria was examined by varying the injected dose and the time of injection in both an albino and a normal (pigmented) strain. Albino nymphs injected with a high dose (100pmol) of [His(7)]-corazonin at the beginning of the third instar turned completely black in the following instar, whereas those injected with the same dose in the middle of the instar developed black patterns with an orange background color, the body coloration characteristic of normal gregarious (crowded) individuals. Injection at the end of the third instar induced a reddish color with few black spots. Irrespective of the time of injection of the peptide, most of these individuals became completely black after ecdysis to the fifth instar. A similar result was obtained with a lower dose (1pmol), although the color expressed was lighter. In the normal strain, injection of 1nmol or 100pmol into crowded third instar nymphs also caused most of them to become completely black in the fourth and fifth instars, but a lower dose apparently had no influence. These results suggest that the temporal changes in hemolymph titer of [His(7)]-corazonin are important in the expression of body color in L. migratoria.     

Phase-related morphological changes induced by [His7]-corazonin in two species of locusts, Schistocerca gregaria and Locusta migratoria (Orthoptera: Acrididae)

The effects of a neurohormone, [His(7)]-corazonin, on phase-related morphological traits (F/C and E/F ratios; F = length of the hind femur, C = maximum width of the head; E = length of fore wing) were re-examined in the desert locust, Schistocerca gregaria Forsk?l. The F/C ratio was significantly different between adults with five and six nymphal instars, respectively, indicating that they need to be analysed separately. Injections of the synthesized peptide (1 nmol) into individually-reared (solitary) nymphs at the second and third instars caused a shift in classical morphometric ratio towards the value typical for crowded (gregarious) individuals in both sexes. The E/F ratio, which is smaller in solitary locusts than in gregarious ones, was also influenced significantly by injections of [His(7)]-corazonin into individually-reared locusts. The effect of [His(7)]-corazonin on E/F ratios was shown more clearly when the nymphs were injected at a higher dose (2 nmol) at the beginning of the third instar. Single injections of the peptide into individually-reared nymphs at different instars revealed that the earlier the injection the larger the 'gregarizing' effects of the peptide on F/C and E/F ratios. The same tendency was also detected in Locusta migratoria Linnaeus. These results supported the hypothesis that [His(7)]-corazonin plays an important role in the control of phase polymorphism in locusts.     

Fine structure of the antennal sensilla of the millipede Orthomorphella pekuensis (Polydesmida: Paradoxosomatidae)

The fine structural characteristics of various sensory receptors on the antenna of a millipede, Orthomorphella pekuensis, were observed with field emission scanning electron microscopy. The antenna of this millipede has eight segments, called articles. On the surface of the antenna, there are a variety of sensory receptors, including olfactory and mechanical receptors. According to their morphological and fine structural characteristics, we could identify four basic types of antennal sensillum: chaetiform sensilla (CS), trichoid sensilla (TS), basiconic sensilla (BS) and apical cone sensilla (AS). The BS are divided further into three subtypes: large basiconic sensilla (BS₁) on the 5th and 6th articles; small basiconic sensilla (BS₂) on the 5th article; and a distinct type of basiconic spiniform sensilla (BS₃) on the 7th article. The most prominent sensilla are four large AS on the distal tip of the 8th segment. Based on our results, we conclude that the main function of the CS and TS are related to mechanical reception, and that the BS and AS are likely to function in olfactory reception of volatile odors of plants, as these sensilla have base and apex pores, respectively.     

苹果榕传粉小蜂雌性触角感器及其生态学意义

利用扫描电子显微镜对苹果榕传粉小蜂Ceratosolen sp.雌性触角感器进行观察。结果显示:苹果榕传粉小蜂雌性触角呈膝形,分布着7类13种类型的感器,包括毛型感器、多孔的板型感器、锥型感器(2种类型)、栓锥型乳突状感器、刺型感器(4种类型)、腔锥型感器(3种类型)和一种专一性的角锥型感器。对各感器的形态、数量、分布进行描述,并结合感器的选择性染色(Ag+染色)、传粉小蜂行为及其榕/蜂互惠共生系统,对不同类型感器的功能和生态学意义进行探讨。     

六斑月瓢虫触角及感觉器扫描电镜观察

[目的] 明确六斑月瓢虫雌雄成虫触角感觉器种类、分布及形态特征。[方法] 利用扫描电子显微镜对六斑月瓢虫雌、雄成虫触角形态及触角感受器超微结构进行观察。[结果] 六斑月瓢虫成虫触角由柄节、梗节和鞭节组成,柄节长度与宽度显著大于梗节长度与宽度;鞭节分为9个亚节,末端3节横向膨大呈锤状。雌雄成虫触角上共有8种感觉器:刺形感觉器(SC)、毛形感觉器(ST)、锥形感觉器(SB)、腔形感觉器(CaS)、钟形感觉器(CS)、哑铃形感觉器(DS)、香肠形感觉器(SS)及B?hm氏鬃毛感觉器(BB)。以毛形感觉器和刺形感觉器分布最广,遍布触角;B?hm氏鬃毛仅存在于触角柄节与梗节;触角鞭节第9亚节顶端密布7种触角感觉器。六斑月瓢虫雌雄成虫触角长度、触角感觉器类型及分布无显著差异。[结论] 六斑月瓢虫成虫触角上共有8种感觉器,其触角可能具有感知机械刺激、识别化学信息素及感受温湿度变化的作用。本研究为进一步了解六斑月瓢虫触角与其行为间的关系提供基础资料。     

Variation in antennal sensory system in different phenotypes of large fruit-tree tortrix Archips podana scop. (Lepidoptera: Tortricidae)

Quantitative composition of antennal sensilla was studied in different phenotypes of large fruit-tree tortrix imago. Changes in the number of sensilla were observed in individuals reared on lilac: male phenotypes AB and B differed by the number of chaetoid, coeloconic, auriculate, and basiconic sensilla; female offspring of males with AB and B phenotypes differed by the number of short trichoid sensilla and auriculate sensilla of the first type. The relationship between male phenotype and the number of sensilla as well as the impact of parental male genotype on the variation in antennal sensory system in female offspring are discussed.     

飞蝗变型及体色多型的内分泌控制机理

飞蝗具变型现象,散居型和群居型在形态特征、生理机能、行为及体色等方面存在明显差异。保幼激素已被证实能诱导飞蝗散居型绿色的出现。近年,从飞蝗心侧体成功地分离了一种神经肽——黑化诱导激素([His^7]-corazonin),并用白化型进行检测,证实了其对体色黑化作用的活性。不同时间对飞蝗若虫注射不同剂量的[His^7]-corazonin,能诱导出现除绿色以外的散居型体色,如浅褐色、褐色、赤褐色、黑色等;也能诱导出现群居型的体色,即黑色配以橘黄色的底色。而且,对散居型若虫注射[His^7]-corazonin能诱导其形态向群居型转换。这些研究证实[His^7]-corazonin对飞蝗的变型有着重要的控制作用,但又不是唯一的。     

The dark-color inducing neuropeptide, [His(7)]-corazonin,causes a shift in morphometic characteristics towards the gregarious phase in isolated-reared (solitarious) Locusta migratoria

The neurohormone, [His⁷]-corazonin is known to induce dark color in the cuticle and epidermis of Locusta migratoria. In the present study, we examined the effects of this hormone on development and morphometrics in two strains, albino and normal, of this locust under isolated conditions. Injection of [His⁷]-corazonin induced dark color in both strains. In either strain, [His⁷]-corazonin injected at the second and third instars did not affect duration of nymphal development or the number of nymphal instars. The shape of the pronotum was more convex in isolated-reared animals than in crowd-reared ones, and injection of [His⁷]-corazonin caused isolated-reared animals to develop a less convex pronotum in the normal strain injected at a high dose (1 nmol×2) but not in the albino strain injected at a low dose (50 pmol×2). [His⁷]-corazonin injected into isolated-reared nymphs caused a shift in 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) towards values typical for crowd-reared (gregarious) individuals of the two strains. This study demonstrated for the first time that [His⁷]-corazonin affected morphometric characteristics in L. migratoria.     

Distribution of olfactory and some other antennal sensilla in the male click beetle Agriotes obscurus L. (Coleoptera : Elateridae)

The distribution of 5 types of sensilla was statistically analysed on the 4–10th antennal segments of the male click beetle Agriotes obscurus (Coleoptera : Elateridae). The distribution pattern of the trichoid pheromone receptors (T₂ sensilla) and the olfactory basiconic B₁B₂ sensilla on the antennae of male A. obscurus differs significantly from the distribution pattern of the contact chemoreceptors (T₁ sensilla) and probably the non-olfactory B₇ and D sensilla. A significant peculiarity of the distribution of olfactory sensilla is their location on the antennal segments as 2 separate (dorsal and ventral) fields of sensilla. The numbers of T₂ and B₁B₂ sensilla on dorsal fields of sensilla of the 4–10th segments increase towards the apex of the antenna nearly linearly. On ventral fields of sensilla of the 4–10th antennal segments, the number of B₁B₂ sensilla is nearly uniform; the number of T₂ sensilla in the proximal part of the antenna increases towards the apex, but on distal segments of the antenna their number stabilizes. It is characteristic of both the T₂ and to B₁B₂ sensilla that their numbers are slightly greater on anterior than posterior sides of dorsal sensillar fields, and also greater on posterior than anterior sides of ventral sensillar fields of all antennal segments investigated. We assume that the number of olfactory sensilla on the antennae of male beetles coincides with the distribution of strength of olfactory signal on the antennae of beetles orientating in an odour plume. The distribution patterns of T₂ and B₁B₂ sensilla of the male A. obscurus can be related to some behavioural peculiarities of olfactory orientation (walking or flying and vibrating of the antennae).     

Corazonin in insects

Corazonin is a peptidergic neurohormone of insects that is expressed in neurosecretory neurons of the pars lateralis of the protocerebrum and transported via nervi corporis cardiaci to the storage lobes of the corpora cardiaca. This peptide occurs with a single isoform in all insects studied so far, with the exception of the Coleoptera in which no corazonin form could be detected. Very few modifications of [Arg(7)]-corazonin, originally isolated from cockroaches, are known, namely [His(7)]-corazonin which is expressed in certain locusts and the stick insect Carausius morosus, and [Thr(4), His(7)]-corazonin recently described from the honey bee Apis mellifera. In this study, we performed a comprehensive screening for corazonin in the different insect groups after detecting of a fourth isoform in a crane fly, Tipula sp. ([Gln(10)]-corazonin). [Arg(7)]-corazonin is distributed in most major lineages of insects, and is thus the ancient form which was present at the time the phylum Insecta evolved. The replacement of Arg with His at position 7 from the N-terminus occurred several times in the evolution of insects. The third isoform, [Thr(4), His(7)]-corazonin, seems to be restricted to bees (Apidae); whereas wasps (Vespidae) and a bumble bee (Apidae) express other corazonins, specifically [His(7)]-corazonin and [Tyr(3), Gln(7), Gln(10)]-corazonin, respectively. A novel corazonin form, [His(4), Gln(7)]-corazonin, was also detected in all South African members of the newly described insect order Mantophasmatodea. The [His(4), Gln(7)]-corazonin separates these species from the Namibian Mantophasmatodea which express [Arg(7)]-corazonin and can be used as a distinct character to distinguish these morphologically similar insects.     

Morphogenesis of the antenna of the male silkmoth. Antheraea polyphemus, III. Development of olfactory sensilla and the properties of hair-forming cells

During adult development of the male silkmoth Antheraea polyphemus, the anlagen of olfactory sensilla arise within the first 2 days post-apolysis in the antennal epidermis (stage 1-3). Approximately on the second day, the primary dendrites as well as the axons grow out from the sensory neurons (stage 4). The trichogen cells start to grow apical processes approximately on the third day, and these hair-forming 'sprouts' reach their definite length around the ninth day (stages 5-6). Then the secretion of cuticle begins, the cuticulin layer having formed on day 10 (stage 7a). The primary dendrites are shed, the inner dendritic segments as well as the thecogen cells retract from the prospective hair bases, and the inner tormogen cells degenerate around days 10/11 (stage 7b). The hair shafts of the basiconic sensilla are completed around days 12/13 (stage 7c), and those of the trichoid sensilla around days 14/15 (stage 7d). The trichogen sprouts retract from the hairs after having finished cuticle formation, and the outer dendritic segments grow out into the hairs: in the basiconic sensilla directly through, and in the trichoid sensilla alongside, the sprouts. The trichogen sprouts contain numerous parallel-running microtubules. Besides their cytoskeletal function, these are most probably involved in the transport of membrane vesicles. During the phase of cuticle deposition, large numbers of vesicles are transported anterogradely from the cell bodies into the sprouts, where they fuse with the apical cell membrane and release their electron-dense contents (most probably cuticle precursors) to the outside. As the cuticle grows in thickness, the surface area of the sprouts is reduced by endocytosis of coated vesicles. When finally the sprouts retract from the completed hairs, the number of endocytotic vesicles is further increased and numerous membrane cisterns seem to be transported retrogradely along the microtubules to the cell bodies. Here the membrane material will most probably be used again in the formation of the sensillum lymph cavities. Thus, the trichogen cells are characterized by an intensive membrane recycling. The sensillum lymph cavities develop between days 16-20 (stage 8), mainly via apical invaginations of the trichogen cells. The imago emerges on day 21.     

Analysis of immunocytochemical staining patterns in the antennal system of Drosophila melanogaster

By immunizing mice with homogenized brains, heads, or a mixture of heads and antennae of D. melanogaster, we obtained six monoclonal antibodies (mabs) that bind to the olfactory system of Drosophila with various degrees of specificity. They can be divided into three groups with respect to their staining pattern: (1) The antibodies ca51/2, na21/2, and nb230 label both in the third (olfactory) antennal segment and in the visual ganglia. All of them bind to antennal structures that can be correlated with basiconic sensilla. The antibody ca51/2 labels sensory neurons of these sensilla. In the antenna of the lozenge ³ mutant, which lacks basiconic sensilla, no labeling is present. In Western blots ca51/2 recognizes in the antenna an antigen of 43.5 kDa, which is expressed in the antenna only in the presence of basiconic sensilla. The antibody na21/2 binds to basiconic and coeloconic sensilla, most likely to the apical part of sheath cells. In immunoblots it recognizes in the antenna two antigens of 42.2 kDa and 46.7 kDa. The latter appears to be correlated in the antenna with the presence of basiconic sensilla. (2) The staining pattern of antibody nc10 is associated with the sheath cells of basiconic and coeloconic sensilla. Moreover, nc10 binds to a subset of glomeruli in the antennal lobe. (3) The staining pattern of the antibodies VG2 and I24B5 is restricted to the antenna. I24B5 recognizes coeloconic sensilla and VG2 recognizes both coeloconic and basiconic sensilla. Staining patterns in both cases include sheath cells.     

Phenotypic plasticity in numbers of antennal chemoreceptors in a grasshopper: effects of food

Grasshoppers, Schistocerca americana, reared from hatching on artificial diet had fewer sensilla on the antennae in the final larval stage than insects reared on lettuce. This was true of basiconic and coeloconic sensilla (presumed olfactory) and trichoid sensilla (presumed gustatory). The degree of difference varied along the antenna and with sensillum type. Adding salicin to the diet restored the numbers of all types of sensillum to levels equal to, or approaching, those in lettuce-fed insects. The addition of some volatile compounds – carvone (monoterpene), chalcone (flavonoid), citral (monoterpene) and guaiacol (phenolic) – resulted in slight increases in number, but coumarin (phenylpropanoid) had no effect. None of the compounds, either singly or in combination, produced more sensilla than were present in plant-fed insects. Accepted: 26 January 1998     

Scanning electron microscopy studies on the first-instar larva of Dermatobia hominis

Abstract. First-instar larvae of Dermatobia hominis collected 1, 4 and 7 days after having penetrated experimentally infected rats, were studied by scanning electron microscope (SEM) observation. On the pseudocephalon there are basiconic and trichoid sensilla (antennal sensory complex), and basiconic, coeloconic and campaniform sensilla (maxillary sensory complex). The thoracic segments bear several rows of small, backwardly pointed, spines, and trichoid, campaniform, coeloconic and pit sensilla. The anterior spiracle is a minute opening. Both small and large spines directed posteriorly are on the first to fourth abdominal segments, which also bear coeloconic and companiform sensilla. These sensilla are present on the unarmed (fifth and sixth) and armed (seventh) abdominal segments. The seventh and the last (eight) abdominal segments have forwardly directed spines. Each spiracular plate has two spiracular openings and four spatulate-like structures called sun rays. The anus and the coeloconic sensilla are proeminent on the last segment. The results are compared with other parasitic dipteran larvae, and emphasize that the multiple types of sensilla on D. hominis larva may have importance in establishing the parasitic phase of the life cycle of this insect.     

伪鞘榕小蜂雌雄成虫触角感器的超微形态、分布及适生意义

伪鞘榕小蜂Sycoscapter trifemmensis是一种寄生于鸡嗉子榕间花期榕果的专性寄生蜂,雌雄两性繁殖策略分化明显,为更好理解和诠释雌蜂寄主定位和雄蜂配偶识别机制,有必要对两性的触角感器进行观察。运用环境扫描电镜观察,对比和探讨了伪鞘榕小蜂雌雄成虫的触角和触角感器类型、分布、数量及其生态适应性。结果表明:雌蜂触角鞭节由11鞭小节组成,总长817.82±33.23μm,分布有毛形感器、刺形感器(类型1)、锥形感器(类型1)、多孔板形感器(类型1)、栓锥形乳突状感器5类5种;雄蜂触角鞭节仅由6鞭小节组成,全长为雌蜂的1/3,且各节有明显的缩短和增粗特征,着生感器包括毛形感器、刺形感器(类型2和类型3)、锥形感器(类型1和类型2)、多孔板形感器(类型2)、腔锥形感器5类7种。雌蜂触角感器的数量与分布显著高于雄蜂,且同类型感器在雌蜂上具有明显的延伸、增粗、分支的特征,以板形感器和锥形感器最为突出。伪鞘榕小蜂雌雄成虫的触角及其感器有明显的性二型,特别是与化学信息识别相关的感器,反映了雌雄蜂在不同生态环境和繁殖压力下的形态分化、行为策略和生态适应。     

Degradation profile of [His7]-corazonin in the hemolymph of the desert locust Schistocerca gregaria

Degradation of the neuropeptide [His7]-corazonin, a key hormone in phase transition in locusts was studied using [3H][His7]-corazonin, RP-HPLC and mass spectrometry. After 4h incubation, 50 and 75% of [His7]-corazonin could still be found in hemolymph of gregarious and solitarious Schistocerca gregaria, respectively. Under in vivo conditions the half-life was 30 min. These results are in contrast to many other neuropeptides that usually have half lives of a few minutes. The peptide is cleaved first by an endopeptidase, either just before or after the Tyr residue at position 5. Next, the C-terminal degradation fragments are further degraded by a dipeptidyl-peptidase, whereas the N-terminal fragments are further broken down one amino acid at a time. In addition, [Dopa5][His7]-corazonin was detected. Upon synthesis, this unexpected molecular modification turned out to be biologically active in bringing about cuticular melanization.     

Sex-specific antennal sensory system in the ant <Emphasis Type="Italic">Camponotus japonicus</Emphasis>: structure and distribution of sensilla on the flagellum

The antennae are a critically important component of the ant’s highly elaborated chemical communication systems. However, our understanding of the organization of the sensory systems on the antennae of ants, from peripheral receptors to central and output systems, is poorly understood. Consequently, we have used scanning electron and confocal laser microscopy to create virtually complete maps of the structure, numbers of sensory neurons, and distribution patterns of all types of external sensilla on the antennal flagellum of all types of colony members of the carpenter ant Camponotus japonicus. Based on the outer cuticular structures, the sensilla have been classified into seven types: coelocapitular, coeloconic, ampullaceal, basiconic, trichoid-I, trichoid-II, and chaetic sensilla. Retrograde staining of antennal nerves has enabled us to count the number of sensory neurons housed in the different types of sensilla: three in a coelocapitular sensillum, three in a coeloconic sensillum, one in an ampullaceal sensillum, over 130 in a basiconic sensillum, 50–60 in a trichoid-I sensillum, and 8–9 in a trichoid-II sensillum. The basiconic sensilla, which are cuticular hydrocarbon-receptive in the ant, are present in workers and unmated queens but absent in males. Coelocapitular sensilla (putatively hygro- and thermoreceptive) have been newly identified in this study. Coelocapitular, coeloconic, and ampullaceal sensilla form clusters and show biased distributions on flagellar segments of antennae in all colony members. The total numbers of sensilla per flagellum are about 9000 in unmated queens, 7500 in workers, and 6000 in males. This is the first report presenting comprehensive sensillar maps of antennae in ants.     

Comparison of antennal sensilla of Anaphes victus and A. listronoti (Hymenoptera, Mymaridae), egg parasitoids of Curculionidae

Antennal sensilla were compared in females and males of two sympatric mymarid Hymenoptera, Anaphes victus and A. listronoti which are, respectively, solitary and gregarious parasitoids of eggs of the carrot weevil Listronotus oregonensis (Coleoptera, Curculionidae). Both species are morphologically very similar in the area where they are sympatric. The external morphology of the sensilla was studied using scanning electron microscopy. Female antennae have seven different types of sensilla, morphologically similar in the two species: trichoid sensilla, which are putative mechanosensilla, sensilla chaetica types 1, 3 and 4, which are presumably contact chemosensilla, and sensilla chaetica type 2 and basiconic and placoid sensilla, which are presumed to be olfactory sensilla. The major difference between the two species is the number of sensilla chaetica type 4, of which 6–9 are found on the antennal club in A. victus, while 10–12 are present in A. listronoti. The antennae of the males of both species are similar in morphology and in the number and distribution of their four types of sensilla, i.e. trichoid sensilla, sensilla chaetica type 1 and basiconic and placoid sensilla. Accepted: 23 November 1998