The paired box gene pox neuro: a determinant of poly-innervated sense organs in Drosophila.

This study describes the structure and function of pox neuro (poxn), a gene previously isolated by virtue of a conserved domain, the paired box, which it shares with the segmentation genes paired and gooseberry. Its expression pattern has been analyzed, particularly during development of the PNS. We propose that poxn is a "neuroblast identity" gene acting in both the PNS and the CNS on the basis of the following evidence. Its expression is restricted to four neuronal precursors in each hemisegment: two neuronal stem cells (neuroblasts) in the CNS, and two sensory mother cells (SMCs) in the PNS. The SMCs that express poxn produce the poly-innervated external sense organs of the larva. In poxn- embryos, poly-innervated sense organs are transformed into mono-innervated. Conversely, ectopic expression of poxn in embryos transformed with a heat-inducible poxn gene can switch mono-innervated to poly-innervated sense organs. Expression of poxn in the wing disc is restricted to the SMCs of the poly-innervated sense organs, suggesting that poxn also determines the lineage of poly-innervated adult sense organs.     

Development of the sensory system in larvae and pupae of Chaoborus crystallinus (DeGeer, 1776; Diptera, Chaoboridae): sensory cells, nerves and ganglia of the tail region

Using various microscopical techniques, we have studied changes in the sensory equipment and architecture of the peripheral nervous system (PNS) around the first metamorphic molt from larva to pupa in the phantom midge Chaoborus. The transparent larvae and pupae of this dipteran with ancestral features allow us to investigate sensilla and their central projections from whole-mount preparations of complete groups of segments. Each sensillum on the posterior larval and pupal segments was identified using its external shape and position, and the morphology of the abdominal ganglia and segmental nerves was investigated. In addition, retrograde fills with the carbocyanine dye DiI were used to trace the axonal paths of most of the extero- and proprioreceptors. These findings were combined to produce maps of the sensory elements of larval and pupal abdomens that were analyzed at three levels: seriality (homonomy), ontogenetic changes of individual sensilla, and homology of the PNS between different species. Comparison of different segments shows for both stages that primarily there is a homonomous basic design of the PNS, but segment-specific modifications are evident in segments 8-10. Comparison of corresponding larval and pupal segments shows that many sensilla retain their internal structure and axonal projections. However, their external cuticular parts are changed in relation to the different life habits of larvae and pupae. Furthermore, some sensilla are completely reduced during the pupal molt, especially those of the tenth segment which appears as a distinct larval structure (caenogenesis). Comparison between species indicates that despite the varying types of sensilla their basic segmental arrangement and their axonal trajectories are conserved.     

Sensory organs of the thoracic legs of the moth Manduca sexta

Summary The thoracic legs of the moth Manduca sexta acquire a new form and develop a new complement of sensory organs and muscles during metamorphosis from larva to adult. Because of our interest in the reorganization of neural circuitry and the acquisition of new behaviors during metamorphosis, we are characterizing sensory elements of larval and adult legs so that we may determine the contribution of new sensory inputs to the changes in behaviors. Here we describe the sensory structures of adult legs using scanning electron microscopy to view the external sensilla and cobalt staining to examine innervation by underlying sensory neurons. We find that, in contrast to larval legs, the adult legs are covered with a diverse array of sensilla. All three pairs of thoracic legs contain scattered, singly innervated scalelike sensilla. Campaniform sensilla occur singly or in clusters near joints. Hair plates, consisting of numerous singly innervated hairs, are also present near joints. Other more specialized sensilla occur on distal leg segments. These include singly innervated spines, two additional classes of singly innervated hairs, and three classes of multiply innervated sensilla. Internal sensory organs include chordotonal organs, subgenual organs, and multipolar joint receptors.     

The formation of the antenna sensory apparatus in some bug (Heteroptera) species in the course of their postembryonic development

In the antenna sensory apparatus of bugs Coreus marginatus, Cimex lectularius, and Rhodnius prolixus sensilla of the four main types are identified: chaetica, trichodea, basiconica, and coeloconica. Chaetoid sensilla are differentiated into two subtypes: sensilla with cogged cuticles and those with smooth ones; trichoid sensilla were divided into long pointed and short ones with blunt tips. In larvae and adults of R. prolixus trichobothria (long filiform hairs) were found on the medial side of pedicellum. The postembryonic changes in the quantitative and qualitative composition of the antenna sensory apparatus were assessed using biometric analysis. The greatest increase of sensory organs was observed upon the nymphal ecdysis from the 5th instar to adult.     

The emergence of sense organs in the wing disc of Drosophila

We have examined the origin of a set of precisely located sense organs in the notum and wing of Drosophila, in transformant flies where lacZ is expressed in the progenitor cells of the sense organs (the sensory mother cells) and in their progeny. Here we describe the temporal pattern of appearance and divisions of the sensory mother cells that will form the eleven macrochaetes and the two trichoid sensilla of the notum, and five campaniform sensilla on the wing blade. The complete pattern of sensory mother cells develops in a strict sequence that extends over most of the third larval instar and the first 10 h after puparium formation. The delay between the onset of lacZ expression and the first differentiative division ranges from 30 h, in the case of the earliest mother cells, to 2 h for the latest mother cells. The first division shows a preferential orientation which is also specific for each sensory mother cell. Up to this stage, there is no marked difference between the three types of mechanosensory organs.     

The precocious commitment of wing Anlagen in Tenebrio molitor revealed by the addition of 20-hydroxyecdysone

An injection of 20-hydroxyecdysone (10 mug per animal) 6-13 days after the moult of the last larval instar of Tenebrio molitor induces the development of prothetelic larvae and larval-pupal intermediates. The state of larval-pupal switchover, or commitment, is only disclosed at the time of injection of the moulting hormone. Prothetelic A and B larvae, with small and medium sized wing Anlagen, undergo another larval or pupal instar. Prothetelic C larvae with bigger Anlagen are unable to moult, but the adult programme is expressed. Ecdysed larval-pupal intermediates give more or less perfect adults, while unecdysed mealworms, imprisoned in their larval cuticle, also expressed the adult programme. The commitment of Tenebrio is not a global switchover because a significant asynchronisation is noted between the development of organs considered. Animal crowding induces a delay in the appearance of wing Anlagen.     

Sensory organs on the body parts of the bed-bug Cimex hemipterus fabricius (Hemiptera : Cimicidae) and the anatomy of its central nervous system

Anatomy of the sensory organs on the prominent body parts of the adult bed-bug Cimex hemipterus (Hemiptera: Cimicidae) and its central nervous system (CNS) was studied by light, transmission, or scanning electron microscopy. The distal tips of antenna and rostrum were found to have rich complements of sensilla. The antenna has both olfactory and gustatory sensilla. Olfactory sensilla project to the antennal lobe organized in the form of glomeruli, while the 2nd component, presumably from gustatory sensilla, projects to the suboesophageal ganglion. The ultrastructure of the sensory pegs on the rostrum of C. hemipterus does not resemble the chemosensilla of adult insects; rather they resemble the larval sensilla of Drosophila melanogaster in the maxillary organ. Earlier we believed this to be a gustatory organ. A few similar sensilla also occur on the antenna, indicating its multimodal role. Amongst the 3 types of sensory hairs located on legs, there are only a few gustatory hairs (7–10 hairs) on the tibia. The pointed and serrate mechanosensory hair types occur in abundance; the serrate type are prominently present on the lateral surface of the legs. On other parts of the body such as the thorax or abdomen, serrate hairs are most abundant. Both the distal segment of antenna and rostrum are invested by 2 nerves, where the axon counts of the 2 antennal nerves are 380 and 425, while each rostral nerve on average has 205 axons. Abundant clusters of microtubules were found in the brain, thoracio-abdominal ganglia, leg-nerves, and the space between muscles and cuticle. These conspicuous microtubule-clusters occur in interaxonal space, mainly glial cells, in the nervous system. In addition, the glial cells have osmiophilic junctions amongst themselves. A novel “hinge and joint” system, which controls the cross-section of the food canal and the salivary duct in an inversely related manner, was found in the rostrum of the bed-bug.     

In-Vivo Imaging of the Drosophila Wing Imaginal Disc over Time: Novel Insights on Growth and Boundary Formation

In developmental biology, the sequence of gene induction and pattern formation is best studied over time as an organism develops. However, in the model system of Drosophila larvae this oftentimes proves difficult due to limitations in imaging capabilities. Using the larval wing imaginal disc, we show that both overall growth, as well as the creation of patterns such as the distinction between the anterior(A) and posterior(P) compartments and the dorsal(D) and ventral(V) compartments can be studied directly by imaging the wing disc as it develops inside a larva. Imaged larvae develop normally, as can be seen by the overall growth curve of the wing disc. Yet, the fact that we can follow the development of individual discs through time provides the opportunity to simultaneously assess individual variability. We for instance find that growth rates can vary greatly over time. In addition, we observe that mechanical forces act on the wing disc within the larva at times when there is an increase in growth rates. Moreover, we observe that A/P boundary formation follows the established sequence and a smooth boundary is present from the first larval instar on. The division of the wing disc into a dorsal and a ventral compartment, on the other hand, develops quite differently. Contrary to expectation, the specification of the dorsal compartment starts with only one or two cells in the second larval instar and a smooth boundary is not formed until the third larval instar.     

Sensory organs of the antennae and mouthparts of beetle larvae (Coleoptera)

The sensilla located on the antennae and maxillary and labial palps of the larvae of 64 beetle species from 22 families were studied using electron microscopy. The larvae of beetles living in different habitats and having different trophic specializations possess a uniform structure of the sensory organs. They are composed of two groups of sensilla on the apical and subapical segments of the antennae, one apical group of sensilla on both maxillary and labial palps, and one or several digitiform sensilla on the lateral surface of the maxillary and, occasionally, labial palp. The external morphology of the sensory organs is adaptive and represents modifications of the initial type. Band-shaped sensilla or placoid sensilla, clearly different from the initial sensory organs, appear in some taxa as rare exceptions, while other groups display either partial reduction of the receptor organs (Gyrinidae) or reduction of the cuticular parts of the sensilla (Cantharidae).     

Regeneration of the tibia and somatotopy of regenerated hair sensilla in Schistocerca gregaria (Forskål)

After injury many arthropods are able to regenerate lost body parts and their innervation. Here, regeneration was studied in the desert locust Schistocerca gregaria after amputation of the midleg tibia and tarsus in the first larval instar. A regenerate was formed first in the third larval instar and it increased in size with each larval moult. The regenerate was always unsegmented and remained much shorter than the intact leg parts. The growth rate was initially rather high and decreased thereafter to that of intact parts. The amputation also influenced the growth rate of proximal leg parts (femur and trochanter) resulting in shortened leg segments. The regenerate carried many sense organs like trichoid sensilla and canal sensilla. The primary mechanosensory neurons of the trichoid sensilla projected somatotopically into the mesothoracic ganglion. A comparison of these projections from intact leg segments and regenerates showed a regrow into the target neuropil areas and a restoration of the somatotopy. Intact sensilla on the injured leg and regenerated sensilla expanded their central projections lateral-medially.     

Postembryonic development of the wing imaginal discs in the female wingless bagworm moth Eumeta variegata (Lepidoptera,Psychidae)

The process of wing disc development and degeneration in the bagworm moth Eumeta variegata was investigated histologically. Morphological differences between two sexes first appear in the penultimate (eighth) larval instar. In the male, wing discs proliferate rapidly in the penultimate larval instar and continue proliferating; a conspicuous peripodial epithelium forms in the last (ninth) larval instar. The hemopoietic organs break down in this stage and disappear completely by the prepupal stage. In the female, in contrast, the wing discs remain as in the previous (seventh) instar, without proliferation of cells inside. No peripodial epithelium forms in the penultimate instar or later. Hemopoietic organs are still attached to the wing discs in the last larval instar and the entire wing discs transform into a plain, thick epidermis in the prepupal period. It is suggested that the hemopoietic organs may prevent the wing discs from developing in E. variegata.     

atonal regulates neurite arborization but does not act as a proneural gene in the Drosophila brain

Drosophila atonal (ato) is the proneural gene of the chordotonal organs (CHOs) in the peripheral nervous system (PNS) and the larval and adult photoreceptor organs. Here, we show that ato is expressed at multiple stages during the development of a lineage of central brain neurons that innervate the optic lobes and are required for eclosion. A novel fate mapping approach shows that ato is expressed in the embryonic precursors of these neurons and that its expression is reactivated in third instar larvae (L3). In contrast to its function in the PNS, ato does not act as a proneural gene in the embryonic brain. Instead, ato performs a novel function, regulating arborization during larval and pupal development by interacting with Notch.     

Larval morphology of Metaphycus flavus and its role in host attachment and larval cannibalism

Metaphycus flavus (Howard) (Hymenoptera: Encyrtidae) is a facultatively gregarious endoparasitoid of soft scales (Hemiptera: Coccidae). When it develops in superparasitised hosts, the larvae often attack and consume brood mates six or more days post oviposition. Under our laboratory conditions (25±1°C and 14 hours of light followed by 18±1°C and ten hours of darkness in 50-70% R.H.), M. flavus eggs hatched three days after oviposition. Measurements of the mandibles and tentorium indicate there are four larval instars, and M. flavus reaches the fourth instar by day six post oviposition, and pupates on day eight. Thus, cannibalism among M. flavus larvae occurs during the fourth instar. During this instar, M. flavus larvae separate from their attachment to the scale cuticle, to which they were tethered by a respiratory structure during the previous three larval instars. Once detached, they are free to move within the scale, which increases the probability of larval encounters and aggressive behaviours. Moreover, the mandibles of the fourth instar are better adapted for fighting than are those of the first three larval instars, since they are larger and more sclerotized. The cranium and mouthparts of M. flavus have four different types of sensory organs, some of which are almost certainly olfactory, an unexpected function for a larva that presumably is surrounded by an aqueous medium where gustatory sensilla would seem to be more appropriate. The cranium also bears two pairs of what appear to be secretory pores.     

Reciprocal transplantation of wing discs between a wing deficient mutant (fl) and wild type of the silkworm, Bombyx mori

The wingless mutant flügellos ( fl ) of the silkworm lacks all four wings. Although wing discs of the fl seem to develop normally until the fourth larval instar, wing morphogenesis stops after the fourth larval ecdysis, probably caused by aberrant expression of an unidentified factor, referred to as fl . To characterize factor fl , the wing discs dissected from the wild-type (WT) and fl larvae were transplanted into other larvae and developmental changes of the discs were examined. When the wing disc from a WT larva was transplanted into another WT larva and allowed to grow until emergence, a small wing appeared that was covered with scales. Thus, the transplanted wing discs can develop autonomously, form scales and evert from adult skin. The WT wing discs transplanted into the fl larvae also developed at a high rate. However, the fl wing discs transplanted into the WT larvae did not develop during the larval to pupal developmental stages. These data suggest that the fl gene product (factor fl) works in the wing disc cells during wing morphogenesis. Its function cannot be complemented by hemolymph in the WT larva. It is also implied that the level of humoral factors and hormones required for wing morphogenesis are normally maintained in the fl larva.     

枣球蜡蚧雄虫不同发育期的形态特征

采用光学显微镜研究了枣球蜡蚧Eulecaniumgiganteum (Shinji)雄虫在不同发育期的形态特征。结果表明 :该虫雄性完成其生活史需经历初孵若虫、固定若虫、预蛹和蛹、雄成虫 4个阶段。初孵的 1龄若虫为扩散活动期 ,虫体具有发达的单眼、触角和 3对胸足 ,便于寻找寄生场所 ,但泌蜡腺体很少。 1龄后期和 2龄若虫为固定取食期 ,虫体缘毛粗大 ,泌蜡腺体很多 ,蜡泌物在虫体背面堆积成龟背状蜡壳 ,自我保护性很强。预蛹和蛹不再取食 ,虫体隐藏在白色的半透明蜡壳内。与若虫相比 ,体形变化很大 ,预蛹出现翅芽 ,而其它各器官退化 ,眼无 ,触角和足短小且分节不显。到蛹期 ,翅芽、触角和足明显长大 ,并出现感觉毛。成虫期为活动交配期 ,头部出现发达的 5对单眼 ,触角 1 0节 ,密布各种刚毛等感觉器。前翅和足发育完好。     

External morphology of antennae and their sense organs in the roach Gromphadorhina brunneri (Blattoidea: Dictyoptera)

The antennae and their sense organs in nymphs and adult roaches of Gromphadorhina brunneri, were investigated and described. The number of segments and sensillae of the nymphal antennae depend on the developmental stage. Sexual dimorphism is pronounced. Males have longer antennae than females as well as an abundance of especially long sensory hairs (long wavy hairs), which are probably responsible for the perception of female sex pheromones. They also have more thin-walled sensory hairs, for instance, sensilla trichodea. On a morphological basis the sensillae of Gromphadorhina brunneri, were named and classified. Long wavy hairs and large sensory hairs appear to be present also in a related species, G. portentosa, but are lacking in others. Their distribution on the antennae varies greatly from that in G. portentosa but their structure varies only slightly. These two types of sense organs are considered to be specialized forms of sensilla chaetica. They are contact chemoreceptors, as are two other types of sensilla chaetica. Furthermore, thin-walled chemoreceptors are present, such as sensilla trichodea, sensilla basiconica, sensilla coeloconica and a typical mechanoreceptor, the sensillum campaniformium.