Abdominal plates,spiracles and sternites in the ventral mesosoma of embryos of the desert scorpion Paruroctonus mesaensis (Scorpiones,Vaejovidae)

Summary

The scanning electron microscope was used to study changes in the ventral mesosoma of the vaejovid scorpion, Paruroctonus mesaensis. Observations are compared with those from scorpion fossils. The oldest fossils are from the Silurian period; migration from water to land occurred in the Carboniferous and Permian periods. All recent scorpions are terrestrial with four pairs of booklungs and spiracles in mesosomal sternites. Ancient eurypterids and scorpions had flap-like abdominal plates attached to the ventral surface of five mesosomal segments. The abdominal plates were apparently an aquatic adaptation, and authors have described possible gill tissue in the chamber above. In scorpion embryos, rectangular (holostem) plate-like structures precede the formation of sternites in the ventral mesosoma. Transverse folds were seen in the space above the abdominal plates. The lack of elaborate gill-like structures here supports an earlier hypothesis that aquatic scorpions had other mesosomal respiratory sites (e.g., pectines), resulting in less reliance on respiratory tissues above the abdominal plates. Spiracles initially appear as round or ovoid patterns in the epidermis at the latero-posterior margins of the ventral plates. The booklung spiracles are positioned farther anterior in sternites, but the developmental sequence for this transition is still unclear and may occur later than the stages of this study. The abdominal plates lengthen and enlarge laterally and/or epidermis is added at the lateral edges so that broad, overlapping sternites eventually cover the ventral surface of the mesosoma.     

Development of segments and appendages in embryos of the desert scorpion Paruroctonus mesaensis (Scorpiones: Vaejovidae)

The scanning electron microscope was used to study the changing features of scorpion embryos from the blastula through early stages in the development of appendages. The earliest scorpion fossils (Silurian period) have structures more advanced than the embryos herein, so the possibility is considered that these embryos still retain and display some features indicative of evolutionary patterns in adult pre-Silurian ancestors. The blastodisc stage is followed by a knob-like germinal center that gives rise to most of the embryo body. The germinal center elongates on the ventral surface of the spherical yolk mass. The broad cephalic lobe is first delineated from the following pedipalpal segment. The limbbuds for the pedipalps and anterior walking legs appear, as additional segments are added at a growth zone at the rear of the embryo body. Initially, in the cephalic lobe there are no limbbuds; then the cheliceral buds emerge from the posterior part of the lobe. The stomodeum appears first in the anterior half of the cephalic lobe, but an oral groove forms and the mouth is displaced posteriorly within the groove. This repositioning allows space anteriorly for invagination (semilunar grooves) of epithelium for the brain and medial eyes. The mouth is directed ventrally in all stages of this study. The widespread chelicerae are initially posterior to the mouth, but later move anterior and dorsal to it. Small limbbud bulges on mesosomal segments disappear later and never become protruding appendages. Metasomal segments are produced free from the yolk surface in a ventral flexure beneath the embryo body. The telson starts as two spherical lobes, but later elongates and tapers distally, not yet developing the sharp sting (aculeus) seen in Silurian and all subsequent scorpions. The walking legs are digitigrade, as in most fossil aquatic scorpions. Segments are delineated in the appendages; the chelicerae and pedipalps are divided distally for chela (claw) formation. Bilateral swellings (limbbuds) on the third abdominal segment become larger than the others, indicating the site of pectine formation. The early fin-like pectines are somewhat posterior in the mesosoma, suggesting ancestral swimming, maneuvering, and balancing for the elongate abdomen. The pectinal surface is initially smooth but later transverse striations increase the surface area as a possible respiratory adaptation. Pectinal teeth (present in Silurian and all subsequent scorpions) and forward movement and merging of anterior abdominal segments are not yet evident in embryos of this study.     

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.     

普通齿蛉幼虫的呼吸系统及呼吸行为

研究了普通齿蛉Neoneuromus ignobilis Navás幼虫的呼吸系统及其呼吸行为。结果表明:普通齿蛉幼虫为全气门式(10对气门)呼吸系统,前中胸、中后胸之间、腹部8节各有1对气门,腹部8节各有气管鳃1对,前6对细短,管状,有较短绒毛,后2对气管鳃较粗长,呈羽毛状。腹部1～7节各有1对毛簇,第8腹节无毛簇。侧纵干气管较粗,4束,自前胸前缘部分成左右2组,每组两根侧纵干气管,向胸腹部延伸,二级气管分别伸达各个气门和毛簇,腹部每节由毛簇处的二级气管分支而来的三级气管相连或延伸至消化道等处。气管鳃中无气管。有毛簇呼吸、气门呼吸和体壁呼吸3种呼吸方式,在水中以毛簇呼吸为主,在陆上进行气门呼吸和体壁呼吸。     

Morphology of the second- and third-instar larvae of Dermatobia hominis by scanning electron microscopy

Larvae of Dermatobia hominis 10–27 days old were collected from experimentally infected rats and their morphology was studied by scanning electron microscopy. The moult from the second to third instar occurs at 18 days, with emergence from the host at 30 days post-infection. The second-instar larvae bear on the pseudocephalon, antennae (coeloconic sensilla), and coeloconic and basicoconic sensilla on the maxillary sensory complex. The thoracic segments bear small backwardly-directed spines anteriorly and ventral trichoid and campaniform sensilla. The first four abdominal segments have small and large backwardly-directed spines that are absent on segments five and six. The seventh and eighth abdominal segments have medium-sized forwardly-directed spines. Abdominal segments are encircled by campaniform sensilla. The terminal end of the eighth abdominal segment bears the anus, prominent anal lobes and two spiracular openings on each spiracular plate. Spiracular plates show a radial sun ray pattern. The rear abdomen also bears an ecdysal aperture, several pores and eight coeloconic sensilla. Although there are slight morphological differences, the spines (predominantly flat and thorn-like) and sensilla (campaniform and coeloconic) of the third-instar larvae show a similar arrangement to that of second-instar larvae. Thoracic trichoid sensilla are not seen in third-instar larvae. A perispiracular gland aperture is situated above each posterior spiracular opening. These morphological features are compared with those of other cuterebrid larvae.     

Morphology of the first instar of Calliphora vicina, Phormia regina and Lucilia illustris (Diptera, Calliphoridae)

Scanning electron microscopy documentation of first instar Calliphora vicina Robineau-Desvoidy, Phormia regina (Meigen) and Lucilia illustris (Meigen) (Diptera: Calliphoridae) is presented for the first time, and the following morphological structures are documented: pseudocephalon; antenna; maxillary palpus; facial mask; labial lobe; thoracic and abdominal spinulation; spiracular field; posterior spiracles, and anal pad. Light microscopy documentation and illustrations are provided for the cephaloskeleton in lateral and ventral views. New diagnostic features are revealed in the configuration of the facial mask, cephaloskeleton and posterior spiracles. The first instar morphology of C. vicina, Ph. regina and L. illustris is discussed in the light of existing knowledge about early instars of blowflies.     

Morphogenesis during molting of the setae in the statocyst sensilla of the mysid shrimp Neomysis integer (Leach, 1814) (crustacea,mysidacea)

The morphogenesis of the setae in the statocyst sensilla of Neomysis integer was studied. Immediately before ecdysis, a new seta lies inverted between the enveloping cells. All of the nine enveloping cells, except the first one, secrete a well defined part of the new seta. The second, third, fourth, and fifth have a trichogen function; the sixth has a trichogen-tormogen function; and the seventh, eighth, and ninth enveloping cells have a tormogen function. It could not be established whether the dendritic sheath is replaced at molt. In the second and third enveloping cells, there is a differential secretion of cuticular material forming the wall of the distal part of the seta. As a consequence, this wall is not homogeneous. The possible role of this heterogeneity in the formation of the gutter-like apical part of the seta is discussed. A mechanism is proposed by which the pore develops at the transition between the midpart and the apical gutter-like part. Before ecdysis, the distal segments of the sensory cells are still connected with the wall of the old seta in the same way as during intermolt. No degeneration is apparent in the distal segments during preparation for the molt. These morphological findings suggest that sensitivity of the sensilla must be maintained until the moment of the ecdysis.     

Pronymphs,hatching, and proboscis assembly in leafhoppers and froghoppers (Hemiptera,Cicadellidae and Aphrophoridae)

Pharate 1st instar nymphs enclosed in the embryonic cuticle, referred to as pronymphs, were studied in a froghopper Aphrophora pectoralis Mats. (Aphrophoridae) and the leafhoppers Oncopsis flavicollis (L.), Populicerus populi (L.), Alebra wahlbergi (Boh.), Igutettix oculatus (Lindb.), and Scenergates viridis (Vilb.) (Cicadellidae). The species vary in the relative length of the pronymphal antennae and details of sculpturing of the cephalic region. No egg bursting structures were observed, except small denticles on the crown region of S. viridis pronymphs. Rudimentary mandibular and maxillary stylets of a pronymph are external, short, tubular appendages containing tips of the corresponding nymphal stylets, whose more basal parts develop inside of the head. Casting off of the embryonic cuticle results in the nymphal stylets being passively pulled out and assuming a close-set parallel orientation. Once the sheaths of unsclerotized cuticle secreted by the peripodial epithelium and enveloping each developing stylet have been cast off with the exuviae, the bare stylets become squeezed and interlocked into a functional bundle. The roles of the maxillary plates, clypeus, labrum, and labium in the stylet bundle assembly are discussed. The process repeats after each molt.     

Microscopic analysis of mechanosensory system monitoring the dynamic claw actions in the tenebrionid beetle Zophobas atratus

Many insects have a pair of claws on each leg. The distribution of mechanoreceptors that monitor claw actions was examined in the tenebrionid beetle Zophobas atratus. Each claw has 25–45 campaniform sensilla (CS) that detect the claw’s deformation due to substrate engagement. Five CS clusters are observed around the end of the 5th tarsomere (Ta5) in a concave, socket-like structure. The 1st cluster, containing 2–5 CS, is embedded in the unguifer to which the claws are articulated. The symmetrical 2nd and 3rd clusters, each containing two CS, are located bilaterally in the ventrolateral grooves of the sidewall of the socket, into which the unguis retractor plate slides. The 4th and 5th clusters, containing 1–2 CS with two hair sensilla, are localized near the ventrolateral ridges of the socket into which the basal portion of the claw is pressed during maximal claw flexion. In addition, Ta5 has a chordotonal organ of six sensory cells to monitor claw extension. These results suggest that the mechanoreceptor system may directly monitor the precise mechanical states of individual claws and provide the central nervous system with the sensory information required for fine feedback control of movements of the pretarsus and other leg segments for locomotion and other purposes.     

Juvenile hormone acts at embryonic molts and induces the nymphal cuticle in the direct-developing cricket

During embryogenesis of hemimetabolous insects, the sesquiterpenoid hormone, juvenile hormone (JH), appears late in embryogenesis coincident with formation of the first nymphal cuticle. We tested the role of embryonic JH by treating cricket embryos with JH III, or the JH-mimic (JHM) pyriproxifen, during early embryogenesis. We found two discrete windows of JH sensitivity. The first occurs during the formation of the first (E1) embryonic cuticle. Treatment with JHM prior to this molt produced small embryos that failed to complete the movements of katatrepsis. Embryos treated after the E1 molt but before the second embryonic (pronymphal) molt completed katatrepsis but then failed to complete dorsal closure and precociously formed nymphal, rather than pronymphal characters. This second sensitivity window was further assessed by treating embryos with low doses of JH III prior to the pronymphal molt. With low doses, mosaic cuticles were formed, bearing features of both the pronymphal and nymphal stages. The nymphal characters varied in their sensitivity to JH III, due at least in part to differences in the timing of their sensitivity windows. Unexpectedly, many of the JH III-treated embryos with mosaic and precocious nymphal cuticles made a second nymphal cuticle and successfully hatched. JH treatment also affected the growth of the embryos. By focusing on the developing limb, we found that the effect of JH upon growth was asymmetric, with distal segments more affected than proximal ones, but this was not reflected in misexpression of Distal-less or Bric-a-brac, which are involved in proximal-distal patterning of the limb.Edited by P. Simpson     

Structure of female genitalia of glassy-winged sharpshooter, Homalodisca coagulata (Say) (Hemiptera: Cicadellidae)

The functional reproductive morphology of the female glassy-winged sharpshooter, Homalodisca coagulata (Say), is described at both light microscopy and scanning electron microscopy levels. The female has nine abdominal segments; the seventh to the ninth abdominal segments are modified for reproduction; the eighth tergite is reduced to two segments, with the ovipositor partially exposed from the modified ninth segment-the pygofer. The pygofer, covered with trichoid and coeloconic sensilla, almost completely encloses the ovipositor, which consists of three pairs of valvulae and two pairs of valvifers. The first and second valvulae function together for oviposition. The first valvulae are located exterior to the second valvulae, both of which bear many trichoid, campaniform, and coeloconic sensilla. The third valvulae, possessing many coeloconic sensilla, envelope the first and second valvulae. Seven major muscles are found to be associated with the ovipositor and the pygofer. The oviposition process is described with respect to the activity of the valvulae and their associated musculature. The female morphology follows the general pattern of cicadellids as a group.     

Post-autotomy claw regrowth and functional recovery in the snapping shrimp Alpheus angulosus

The ability to regenerate lost tissues, organs or whole body parts is widespread across animal taxa; in some animals, regeneration includes transforming a remaining structure to replace the one that was lost. The transformation of one limb into another involves considerable plasticity in morphology, physiology and behavior, and snapping shrimp offer excellent opportunities for studying this process. We examined the changes required for the transformation of the small pincer to a mature snapping claw in Alpheus angulosus. First molt claws differ from mature claws in overall shape as well as in morphology related to snapping function; nonetheless, shrimp with first molt claws do produce snaps. While most shape variables of second molt claws do not differ significantly from mature claws, the plunger (structure required for snap production) does not reach mature size until the third molt for females, or later for males. Thus, the pincer claw can be transformed into a functional snapping claw in one molt, although both the underlying morphology and superficial shape are not fully regenerated at this stage. The rapid production of a functional snapping claw that we observe in this study suggests that this particular function is of significant importance to snapping shrimp behavior and survival.     

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.     

Respiratory significance of the thoracic and abdominal morphology of Belostoma and Ranatra (insecta,heteroptera)

Summary Both Belostoma and Ranatra possess I–II, subepimeral, thoracic subalar, and abdominal subalar air stores. In Belostoma, unlike Ranatra, the subepimeral air store is greatly enlarged, the abdominal subalar store is partially exposed to the water, and a fully exposed ventral abdominal air store is also present. All the air stores of Ranatra are normally concealed.The mesothoracic and metathoracic spiracles, which open onto the I–II and subepimeral air stores respectively, are of limited permeability. They appear to have less respiratory importance than the large and highly permeable first abdominal spiracles, which lie in the subalar air space and can probably exhale and inhale large amounts of air. The large eighth abdominal spiracles, which lie at the base of the siphon or retractile organ, can also inhale or exhale much air in Ranatra but appear to be mainly exhalant in Belostoma. The smaller second through seventh abdominal spiracles structurally resemble the eighth ones in Belostoma and open onto the ventral abdominal air store. In Ranatra they appear to have no significant respiratory function.Both genera obtain atmospheric air and give off exhaled air by means of the posterior retractile organ or siphon. The two types of air appear to follow different pathways in the two genera. In Ranatra atmospheric air appears to enter the tracheal system mainly or entirely through the eighth abdominal spiracles and then passes through the first abdominal spiracles into the subalar space. Exhaled air follows the reverse pathway. In Belostoma, however, atmospheric air probably enters the tracheae mainly through the first abdominal spiracles; it is conveyed to these spiracles from the retractile organ through the subalar space or, more indirectly, through the ventral abdominal air store. Air exhaled through the first abdominal spiracles follows the reverse route; the eighth abdominal spiracles can also exhale directly into the base of the retractile organ.During underwater respiration the abdominal portion of the subalar air store appears to be the main reservoir for oxygen. The subalar oxygen is initially atmospheric, and is supplemented, during submersion, by other sources of oxygen. Belostoma may use its exposed ventral abdominal air store, and its partially exposed abdominal subalar one, as physical gills; both these stores communicate with the inhalant first abdominal spiracles. Ranatra, none of whose air stores are normally exposed, appears, to be less capable of utilizing dissolved oxygen, but the considerable amount of atmospheric oxygen in the elongated siphon may be inhaled, during submersion, through the eighth abdominal spiracles.In both genera the thoracic air stores appear to be of less respiratory importance than the abdominal ones. They do not appear capable of obtaining large amounts of oxygen, and the thoracic spiracles are relatively impermeable. All the air stores, however, serve to protect the spiracles against the entry of water, and also contribute to the body's hydrostatic balance. It is also possible that some of the air stores play a role in pressure reception.The literature indicates much intergeneric variation in the respiration of Belostomatidae and Nepidae. In the Belostomatidae there is considerable variation in the extent of the ventral abdominal air store and in the roles of the subalar air store and the spiracles. The Nepidae show differences in their ability to utilize dissolved oxygen and in the extent of the subepimeral air store.     

Development of phenotypic differences in sensillum populations on the antennae of a grasshopper,Schistocerca americana

The development of diet-induced phenotypic differences in numbers of sensilla on the antennae of the grasshopper Schistocerca americana was studied using the exuviae produced at each molt. This made it possible to follow changes within an individual insect. In the first instar, insects had similar numbers of four sensillum types: uniporous trichoid sensilla, coeloconic sensilla, and large and small multiporous basiconic sensilla. Rearing on lettuce resulted in sixth instars with greater numbers of three sensillum types than siblings reared on an artificial diet. The first statistically significant differences between treatments in numbers of trichoid sensilla and large basiconic sensilla occurred in the third and fourth instars, respectively. No major reductions in sensillum numbers occurred at any time and the phenotypic differences resulted from differences in the numbers added at each molt.     

Ecdysis in scorpions: supine behavior and exuvial ultrastructure

Abstract. Ecdysis in scorpions has some common features in all species in which it has been examined. Immature scorpions about to molt become less active, and the cuticle changes in appearance. When humidity and other conditions are suitable, the animal begins ecdysis with cheliceral and pedipalpal movements and internal processes that tear the pleural membrane just ventral to the anterior and lateral edges of the carapace. The carapace is pushed upward from within, and the animal starts to emerge through the opening made by the elevated carapace. This is usually done in the prone position. As the anterior body emerges, the pedipalps and walking legs are stretched posteriorly with their distal ends temporarily confined within the old exoskeleton. The buthid scorpions Androctonus australis and Parabuthus transvaalicus become supine before they begin to emerge. Ecdysis is completed lying supine, and the animal rights itself shortly thereafter. A turnover from upright to supine is not seen at any other time in scorpions. After prone or supine extraction, the exuvium includes the cuticle of booklungs, bristles, and sensilla and is relatively intact except for the wedge-shaped opening at the anterior end.     

On the pronymphal stage in the migratory locust (Locusta migratoria, Orthoptera, Acrididae)

The structure of the integument, somatic and visceral muscles, midgut, and Malpighian tubules were investigated at the late stages of the embryonic and early postembryonic development of the migratory locust, Locusta migratoria, to assess the organization of its pronymphal stage. In its morphogenetic features, the vermiform locust larva sometimes called the pronymph corresponds to the first nymphal instar covered with the second embryonic cuticle which has not been shed. Since the first-instar locust nymphs before and after the shedding of this embryonic cuticle differ significantly in many morphological characters, two consecutive phases of this nymphal instar can be distinguished: the first phase existing from the moment of development of the third embryonic cuticle to the shedding of the second one; the second phase existing from the shedding of the second embryonic cuticle to the formation of the cuticle of the second nymphal instar. Since the pronymphal stage should precede the nymph stage, it may be concluded that the pronymph of the locust is fully embryonized and covered with the second embryonic cuticle, which is also typical of other insects with hemimetabolous development (Konopová and Zrzavý, 2005). Therefore, it would be erroneous to refer to the vermiform first-instar nymph as the pronymph, because the two stages are separated by molting and formation of a new cuticle.     

The ultrastructure of the oenocytes in the molt/intermolt cycle of an insect

The structure and development of the permanent oenocytes of Calpodes ethlius (Lepidoptera, Hesperiidae) are described. There are three sorts of oenocyte. The permanent oenocytes are arranged ventral to the last two pairs of spiracles on abdominal segments 7 and 8 in four clusters of about 45 cells each. The molt cycle oenocytes are ventral to the other spiracles and only enlarge at molting. The subdermal oenocytes differentiate from the epidermis in large numbers shortly before pupation. The permanent oenocytes are large polyploid cells characterized by a cytoplasm of densely packed smooth tubular endoplasmic reticulum, and a plasma membrane invaginated in a meshwork of tubes ending in a reticular layer about 12 micro below the surface. There are two sorts of Golgi complex, one small and of conventional form, the other composed of clouds of microvesicles. 'Dense bodies', believed to belong to the microbody class of organelles, arise directly from the STER. There is a variety of membranous and 'crystalline' inclusions. The formation of isolation membranes from the tubular endoplasmic reticulum, and the origin of isolation bodies and autophagic vacuoles are described. Some autophagy takes place at all times in the molt/intermolt cycle, but there are phases of massive autophagy before the 4th-5th molt and the 5th-pupal molt. These phases coincide with pinocytosis of blood proteins and overlap with or are followed by phases of nuclear replication, RNA synthesis (ribosomes) and ER regeneration. Nuclear blebbing occurs before pupation. The morphology of the oenocytes is most like that of vertebrate cells engaged in steroid hormone synthesis. It is pointed out that the oenocytes rather than the prothoracic glands could be the source of ecdysone and the stimulus for molting.