Sense organs on the antennal flagellum of psocids (Insecta,Psocoptera)

The sense organs on the antennal flagella of five species of winged psocids belonging to two families of Psocoptera, Psocidae and Leptopsocidae, have been examined. All agree in possessing tactile hairs, thick-walled chemoreceptors and long, porous chemoreceptors. Thin-walled chemoreceptors were identified in all species except Metylophorous novaescotiae. Coeloconic chemoreceptors were present in all species except Echmepteryx hageni. Campaniform sense organs were found only in Metylophorus novaescotiae and Psocus leidyi.     

Sense organs on the antennal flagellum of two species of embioptera (Insecta)

Five different kinds of sense organs have been identified on the antennal flagellum of males of Ptilocerembia sp: (1) tactile hairs, (2) thick-walled chemoreceptors, (3) chemoreceptors with thick walls and many pores, (4) thin-walled chemoreceptors and (5) campaniform sense organs. The third type has not been reported previously for other species of insects and combines the characteristics of typical thick-walled and thin-walled chemoreceptors. The campaniform sense organs are situated in sets of three, 120° apart, on the periphery of the distal end of nearly every subsegment. They lie at the transition zone where the thicker cuticle of the subsegment changes to the thin membrane between subsegments.     

Sense organs on the antennal flagella of earwigs (Dermaptera) with special reference to those of Forficula auricularia

Five types of sense organs have been found on the antennal flagellum of Forficula auricularia Linnaeus: (1) tactile hairs; (2) long, thick-walled chemoreceptors; (3) short, thick-walled chemoreceptors; (4) thin-walled chemoreceptors and (5) coeloconic chemoreceptors. Earlier workers have placed the first three of these in a single class. The structural characteristics and distribution of these five kinds of sensilla on the antenna are described. All of the types except the third have been reported previously for other species but short, thick-walled chemoreceptors have not been described before. They are remarkable for the presence of (1) a broad pad-Like structure that absorbs dyes readily at the distal end of the peg and (2) a spheroid body, over a micron in diameter, on the dendrite of each olfactory neuron at the point where it narrows to assume the structure of a cilium. The antennal sense organs of a few specimens of Chelisoches morio (Fabricius), Labidura riparia (Pallas) and Euborellia annulipes (Lucas) were also examined.     

Sense organs on the antennal flagellum of a giant cockroach, Gromphadorhina portentosa, and a comparison with those of several other species (Dictyoptera, Blattaria)

Two kinds of tactile hairs, plain and wavy, thick-walled chemoreceptors and two types of thin-walled chemoreceptors are present on the antennal flagellum of Gromphadorhina portentosa males. These are described and their location on the antenna noted. Females of this species have plain tactile hairs and the same types of chemoreceptors as do the males but wavy tactile hairs are absent. The antennal sense organs of a few specimens of five other species of cockroaches — Periplaneta americana, Blatta orientalis, Supella longipalpa, Pycnoscelus surinamensis and Diploptera punctata — were also examined. All lacked tactile hairs but were provided with thick-walled chemoreceptors and with two types of thin-walled chemoreceptors similar to those described for Gromphadorhina portentosa.     

Sense organs of a thysanuran, Ctenolepisma lineata pilifera, with special reference to those on the antennal flagellum (Thysanura, Lepismatidae)

Six types of sense organs are present on the antennal flagellum of Ctenolepisma lineata pilifera: tactile hairs, trichobothria, thick-walled chemoreceptors, small thin-walled chemoreceptors and coeloconic chemoreceptors. The number, size and distribution on the antenna of each type have been recorded. The base of the tactile hair is more complex than is that of other insects examined earlier. Trichobothria, long, slender hairs that oscillate in a gentle puff of air, are an unusal feature in insects and especially so for the antenna. The two types of thin-walled chemoreceptors differ in shape, size and in the structure of their walls and internal parts. A pocket-like depression of the floor of the cavity in which the peg of the coeloconic sense organ is set has not been found in earlier studies. Its function is unknown. The axons from the sensory neurons extend along the inner surface of the antennal epidermis as a sheet of fibers lining the antennal lumen. Near the pedicel the axons leave the epidermis and join to form the antennal nerve. A few observations on sense organs on appendages other than the antennae and some notes on behavior are included.     

Sensilla on the antennal flagellum of the sawyer beetles Monochamus notatus (Drury) and Monochamus scutellatus (Say) (Coleoptera: Cerambycidae)

The antennae of the sawyer beetles Monochamus notatus and M. scutellatus were examined with the light and scanning electron microscopes to determine the types, number, distribution and innervation of the sense organs. Nine types of sensilla are described. Both short, thin-walled pegs (sensilla basiconica) and reversely curved thick-walled hairs (sensilla trichodea) are chemoreceptors. There are three types of long, thick-walled hairs (sensilla chaetica) which may be mechanoreceptors. One of these is modified in males to form unique snail-shaped pegs. A few dome-shaped organs, probably campaniform sensilla, were found. In addition to sense organs, many glands occur in association with the sensilla, and the antennae are well supplied with dermal glands connected by canals to small pores on the surface.     

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.     

Sense Organs of Two Marine Arthrotardigrades (Heterotardigrada,Tardigrada)

The sense organs of the marine arthrotardigrades Halechiniscus greveni Renaud-Mornant & Deroux and Batillipes noerrevangi Kristensen were investigated with the aid of interference phase-contrast and transmission electron microscopy. The sense organs of the two species are quite different in outer cuticular morphology, but the inner cellular parts are constructed after the same model: the arthropod sensory setae. All investigated sensilla contained ciliary structures. Regeneration of the cirri during moult is very similar to the regeneration process in insects and spiders, but in the two arthrotardigrades the trichogen cell is not retracted from the cuticular part of the setae after moult. The clava is presumed to be olfactory; the cephalic cirri and the leg spines are contact chemoreceptors with one mechanoreceptive cilium terminating at the sensillum shaft. Cirrus E is compared with the trichobothrium of arthropods and the phylogenetic implications of the investigated structures are discussed.     

The nature and development of sex attractant specificity in cockroaches of the genus Periplaneta. I. Sexual dimorphism in the distribution of antennal sense organs in five species

Sexual dimorphism in the distribution of antennal sense organs is common among adults of the genus Periplaneta. In three out of the four strains of Periplaneta americana examined, adult males had more contact chemoreceptors than females. In the fourth strain of P. americana and in P. australasiae, P. brunnea, P. fuliginosa, and P. japonica, no statistically supportable sexual dimorphism of contact chemoreceptors was found. However, in all strains and species of Periplaneta examined, sexual dimorphism was found in the total number and/or density of olfactory sensilla. Male adults had nearly twice as many olfactory sensilla as female adults. These observations are consistent with the behavioral observation that males within the genus Periplaneta rely on the reception of an airborne pheromone for the initiation of courtship behavior. In P. americana, where sexual dimorphism was found in the contact chemoreceptors, contact stimuli release the full wing raising display and presentation in males during courtship.     

Structures on the antennal flagellum of a caddisfly,frenesia missa (Trichoptera,Limnephilidae)

Non-innervated macrotrichia and microtrichia, thick-walled chemoreceptors and three kinds of thin-walled chemoreceptors are present on the antennal flagellum of Frenesia missa. One of the thin-walled receptors, the plate organ, is of a type not previously recorded for any insect. About four times as many plate organs are present on the flagellum of the male as on that of the female. They occur also on the maxillary and labial palps.     

Rôle of gustation and olfaction in food plant discrimination in the tobacco hornworm, Manduca sexta

The rôle of the various chemoreceptors in behavioural discrimination among closely related food plants was studied in Manduca sexta larvae. Amputation of the three types of receptors in various combinations showed that: (1) removal of the maxillary styloconica (gustatory) causes a drastic loss of discrimination; (2) ablation of either the maxillary palpi or antennae (olfactory) also reduces discrimination, but to a lesser degree; and (3) amputation of both palpi and antennae simultaneously causes a severe discriminatory loss comparable to that of gustatory loss. We conclude that both gustation and olfaction are important for host plant discrimination. These chemosensory organs were also shown to play a strong role in the induction of preference. Unilateral extirpations of all three sense organs caused no detectable loss, demonstrating their redundancy in normal animals.     

The organization of the malleolar sensory system in the solpugid, Chanbria sp

Five malleoli extend ventrally from the proximal segments of each hind leg of mature and immature solpugids. Each malleolus contains the dendrites of 72,000 bipolar sensory neurons with somata in a ganglion in the leg just dorsal to the malleolus. The dendrites terminate in a sensory groove which extends along the ventral edge of the sense organ. The sensory groove has a slit-like opening to the outside, suggesting that these sense organs are chemoreceptors. The malleoli also contain long processes from microvillar cells with somata in the sensory ganglion. The microvillar cell processes extend into the malleolus to the ciliary ending of the dendrites where each process has an enlarged microvillar ending which surrounds the outer segments of 15 to 22 dendrites and releases a sense secretion.     

The relationship between O2 chemoreceptors, cardio-respiratory reflex and hypoxia tolerance in the neotropical fish Hoplias lacerdae

The localization, distribution and orientation of O₂ chemoreceptors associated with the control of cardio-respiratory responses were investigated in the neotropical, Hoplias lacerdae. Selective denervation of the cranial nerves (IX and X) was combined with chemical stimulation (NaCN) to characterize the gill O₂ chemoreceptors, and the fish were then exposed to gradual hypoxia to examine the extent of each cardio-respiratory response. Changes in heart rate (f_H) and ventilation amplitude (V_amp) were allied with chemoreceptors distributed on both internal and external surfaces of all gill arches, while ventilation rate (f_R) was allied to the O₂ chemoreceptors located only in the internal surface of the first gill arch. H. lacerdae exposed to gradual hypoxia produced a marked bradycardia (45%) and 50% increase in V_amp, but only a relatively small change in f_R (32%). Thus, the low f_R response yet high V_amp were in accord with the characterization of the O₂ chemoreceptors. Comparing these results from H. lacerdae with hypoxia-tolerant species revealed a relationship existent between general oxygenation of the individual species environment, its cardio-respiratory response to hypoxia and the characterization of O₂ chemoreceptors.     

Internal Sense of Direction: Sensing and Signaling from Cytoplasmic Chemoreceptors

SUMMARY

Chemoreceptors sense environmental signals and drive chemotactic responses in Bacteria and Archaea. There are two main classes of chemoreceptors: integral inner membrane and soluble cytoplasmic proteins. The latter were identified more recently than integral membrane chemoreceptors and have been studied much less thoroughly. These cytoplasmic chemoreceptors are the subject of this review. Our analysis determined that 14% of bacterial and 43% of archaeal chemoreceptors are cytoplasmic, based on currently sequenced genomes. Cytoplasmic chemoreceptors appear to share the same key structural features as integral membrane chemoreceptors, including the formations of homodimers, trimers of dimers, and 12-nm hexagonal arrays within the cell. Cytoplasmic chemoreceptors exhibit varied subcellular locations, with some localizing to the poles and others appearing both cytoplasmic and polar. Some cytoplasmic chemoreceptors adopt more exotic locations, including the formations of exclusively internal clusters or moving dynamic clusters that coalesce at points of contact with other cells. Cytoplasmic chemoreceptors presumably sense signals within the cytoplasm and bear diverse signal input domains that are mostly N terminal to the domain that defines chemoreceptors, the so-called MA domain. Similar to the case for transmembrane receptors, our analysis suggests that the most common signal input domain is the PAS (Per-Arnt-Sim) domain, but a variety of other N-terminal domains exist. It is also common, however, for cytoplasmic chemoreceptors to have C-terminal domains that may function for signal input. The most common of these is the recently identified chemoreceptor zinc binding (CZB) domain, found in 8% of all cytoplasmic chemoreceptors. The widespread nature and diverse signal input domains suggest that these chemoreceptors can monitor a variety of cytoplasmically based signals, most of which remain to be determined.     

Bacterial chemoreceptors of different length classes signal independently

Bacterial chemoreceptors sense environmental stimuli and govern cell movement by transmitting the information to the flagellar motors. The highly conserved cytoplasmic domain of chemoreceptors consists in an alpha‐helical hairpin that forms in the homodimer a coiled‐coil four‐helix bundle. Several classes of chemoreceptors that differ in the length of the coiled‐coil structure were characterized. Many bacterial species code for chemoreceptors that belong to different classes, but how these receptors are organized and function in the same cell remains an open question. E. coli cells normally code for single class chemoreceptors that form extended arrays based on trimers of dimers interconnected by the coupling protein CheW and the kinase CheA. This structure promotes effective coupling between the different receptors in the modulation of the kinase activity. In this work, we engineered functional derivatives of the Tsr chemoreceptor of E. coli that mimic receptors whose cytoplasmic domain is longer by two heptads. We found that these long Tsr receptors did not efficiently mix with the native receptors and appeared to function independently. Our results suggest that the assembly of membrane‐bound receptors of different specificities into mixed clusters is dictated by the length‐class to which the receptors belong, ensuring cooperative function only between receptors of the same class.     

Sense organs on the antennae of two species of thrips (Thysanoptera, insecta)

The antennae of two species of thrips, Bagnalliella yuccae (Hinds) and Frankliniella tritici (Fitch), have been examined with the light and electron microscopes. The antennal flagellum of both species is provided with tactile hairs, thick-walled chemoreceptors and thin-walled chemoreceptors. In addition, B. yuccae, but not F. tritici, has a single coeloconic chemoreceptor on the dorsal surface of the pedicel. Observations were made on the fluids in the lumen of the antennae of E. yuccae in the living insect. The movement of the fluids probably has an important physiological significance.     

Three unrelated chemoreceptors provide Pectobacterium atrosepticum with a broad-spectrum amino acid sensing capability

Amino acids are important nutrients and also serve as signals for diverse signal transduction pathways. Bacteria use chemoreceptors to recognize amino acid attractants and to navigate their gradients. In Escherichia coli two likely paralogous chemoreceptors Tsr and Tar detect 9 amino acids, whereas in Pseudomonas aeruginosa the paralogous chemoreceptors PctA, PctB and PctC detect 18 amino acids. Here, we show that the phytobacterium Pectobacterium atrosepticum uses the three non-homologous chemoreceptors PacA, PacB and PacC to detect 19 proteinogenic and several non-proteinogenic amino acids. PacB recognizes 18 proteinogenic amino acids as well as 8 non-proteinogenic amino acids. PacB has a ligand preference for the three branched chain amino acids L-leucine, L-valine and L-isoleucine. PacA detects L-proline next to several quaternary amines. The third chemoreceptor, PacC, is an ortholog of E. coli Tsr and the only one of the 36 P. atrosepticum chemoreceptors that is encoded in the cluster of chemosensory pathway genes. Surprisingly, in contrast to Tsr, which primarily senses serine, PacC recognizes aspartate as the major chemoeffector but not serine. Our results demonstrate that bacteria use various strategies to sense a wide range of amino acids and that it takes more than one chemoreceptor to achieve this goal.     

Ultrastructural aspects of mechano- and chemoreceptors in Branchiobdella pentodonta (annelida,oligochaeta)

The surface receptors in Branchiobdella pentodonta consist of “sense buttons” prevalent on the prostomium, isolated sense cells all along the body of the animal, and free nerve endings. The “sense buttons” are uni- and multiciliated neurosensitive elements and supporting cells together with mucus glandular processes and muscle fibers. In the neurosensitive elements the cilia are always surrounded by cytoplasmic extroversion. The cytoplasm of the apical zone has abundant small dense granules, mitochondria, bands of tonofilaments, and microtubules. The cilium of uniciliated elements originates from three short roots. The highly vacuolated support cells surround the neurosensitive elements, separating them from each other. The “sense buttons” appear to be mechanoreceptors and chemoreceptors, and the isolated sense cells tactile mechanoreceptors, as are the free nerve endings. The surface receptors are compared with those of other Oligochaeta and Hirudinea.     

Fine structure of tarsal sensory organs in the whip spider Admetus pumilio (Amblypygi, Arachnida).

The sensory organs on the tarsi of the antenniform first legs of the whip spider Admetus pumilio C. L. Koch (Amblypygi, Arachnida) were examined with the scanning and transmission electron microscope. At least four different types of hair sensilla were found: (1) thick-walled bristles, which have the characteristics of contact chemoreceptors (several chemoreceptive dendrites in the lumen plus two mechanoreceptors at the base); (2) short club sensilla, innervated by 4-6 neurons which terminate in a pore on the tip; they are possibly humidity receptors; (3) porous sensilla, which are either innervated by 20-25 neurons and have typical pore tubules, or they have 40-45 neurons but no pore tubules; both types are considered to be olfactory; (4) rod sensilla occur in clusters near segmental borders; they are innervated by only one large dendrite which branches inside the lumen. Other tarsal receptors are the claws, which correspond to contact chemoreceptors, and the pit organ which resembles the tarsal organ of spiders. Compared to other arthropod sensilla, the contact chemoreceptors are very similar to those of spiders, while the porous sensilla correspond structurally to olfactory receptors in insects; the club and rod sensilla seem to be typical for amblypygids.