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.     

Neuroanatomical and electrophysiological studies of identified contact chemoreceptors on the ventral ovipositor valve of 3rd instar larvae of lubber grasshoppers (Taeniopoda eques)

A large number of contact chemoreceptors are located on the ovipositor valves of adult female grasshoppers. These receptors play an important role in many aspects of grasshopper life such as detecting the chemical composition of the soil before and during oviposition. It is surprising, however, to find these types of receptors on the ovipositor valves of instar larvae which are not able to oviposit. Thus, these receptors may serve functions other than to search for a suitable site for egg laying. Observation under the scanning electron microscope revealed the presence of uniporous basiconic contact chemoreceptors in addition to different types of trichoid mechanoreceptors on the ovipositor valve of lubber grasshopper 3rd instar larvae. Neuroanatomical studies have shown that these sensilla are multiply innervated, containing one mechanosensory neuron and four chemosensory neurons that project locally and intersegmentally. The tip recording technique from single basiconic sensilla demonstrated mechanosensory responses to deflections of the sensillum as well as gustatory activity when in contact with different chemical solutions. The electrophysiological studies have shown that these sensilla serve as contact chemoreceptors and not as olfactory receptors.     

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.     

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.     

Preliminary electrophysiological study of the contact chemoreceptors in a spider

During the first stages of their life, young spiders of the species Tegenaria atrica, show behavioural changes that lead them from eusocial groups (5–20 days old) to a solitary life (40 days old). This change is paralleled by an increase in their number of contact chemoreceptors, previously shown to be used during social interactions. We have examined whether these chemoreceptors exhibit functional changes using morphological and electrophysiological techniques. Our results indicate that such changes do not occur at the level of the chemoreceptors, thus suggesting that this behavioural shift has another physiological ground.     

Chemoreceptors in the cibario-pharyngeal pump of the cabbage looper moth,Trichoplusia ni (Lepidoptera: Noctuidae)

The structure of contact chemoreceptors in the cibariopharyngeal pump of the moth Trichoplusia ni (Lepidoptera: Noctuidae) is described. Two types of receptors designated A and B are located on the floor of the pump. Two groups of 9-12 A receptors are located in the anterior part of the pump, and two groups of two B receptors are in the posterior part of the pump. Five sensory dendrites extend to the tip of each A receptor and four to each B receptors. Available evidence indicates that these receptors are contact chemoreceptors and do not serve as mechanoreceptors. The receptors are compared to those of other insects.     

Localization and electrical activity of the distance chemoreceptors that mediate predator avoidance behaviour in Acmaea limatula and Acmaea scutum (gastropoda, prosobranchia).

1. The marine gastropods Acmaea (Collisella) limatula and Acmaea (Notoacmea) scutum respond to distant predatory starfish (i.e. to starfish scent) by moving up a vertical surface. 2. The distance chemoreceptors that mediate this avoidance behaviour are located on the mantle margin. Heat cauterization of the limpets' mantle margin eliminates their responsiveness to Pisaster ochraceus scent, while a similar cauterization of the ctenidium and the osphradia does not diminish the avoidance behaviour. 3. Primary afferent electrical activity can be recorded from the chemoreceptors on the mantle margin that are responsive to starfish scent and also from other physiologically distinct receptors that are responsive to contact with starfish tube feet.     

Structures on the antennal flagellum of a katydid, Neoconocephalus ensiger (Orthoptera, Tettigoniidae)

The long antennal flagellum of Neoconocephalus ensiger is covered with many sharp-tipped hairs that appear to be non-innervated; thick-walled chemoreceptors, that may also have a tactile function; thin-walled chemoreceptors of several kinds and coeloconic chemoreceptors. All of the chemoreceptors are innervated by small groups of neurons. The first flagellar subsegment is unusual in that it bears a small protuberance on its latero-ventral surface. This marks the site of the attachment, internally, of a scoloparium containing about eleven scolopales in which the dendrites of some 23 sensory neurons terminate. The most distal subsegment lacks the scoloparium reported earlier for the grasshopper. No conspicuous difference between the antennae of males and of females was found.     

Alleged silk spigots on tarantula feet: Electron microscopy reveals sensory innervation,no silk

Several studies on tarantulas have claimed that their tarsi could secrete fine silk threads which would provide additional safety lines for maintaining a secure foot-hold on smooth vertical surfaces. This interpretation was seriously questioned by behavioral experiments, and more recently morphological evidence indicated that the alleged spigots (“ribbed hairs”) were not secretory but most likely sensory hairs (chemoreceptors). However, since fine structural studies were lacking, the sensory nature was not proven convincingly. By using transmission electron microscopy we here present clear evidence that these “ribbed hairs” contain many dendrites inside the hair lumen – as is the case in the well-known contact chemoreceptors of spiders and insects. For comparison, we also studied the fine structure of regular silk spigots on the spinnerets and found them distinctly different from sensory hairs. Finally, histological studies of a tarantula tarsus did not reveal any silk glands, which, by contrast, are easily found within the spinnerets. In conclusion, the alleged presence of silk spigots on tarantula feet is refuted.     

Contact chemoreception related to host selection and oviposition behaviour in the monarch butterfly, Danaus plexippus

Abstract. Behavioural events during host selection by ovipositing monarch butterflies (Danaus plexippus (L.), Danainae, Nymphalidae) include tapping the leaf surface with fore-tarsi and touching this surface with mid-tarsi (‘drumming’) and antennae. Flavonoids identified from host plant extracts are known to stimulate oviposition. Scanning electron microscopy revealed the presence of contact-chemoreceptor sensilla on all appendages that contact the leaf surface. This electrophysiological study was conducted to identify the contact chemoreceptors that are sensitive to the known oviposition stimuli and are therefore probably involved in host recognition. Receptor cells of conspicuous sensilla grouped in clusters on fore-tarsi of females were sensitive to the behaviourally active butanol fraction of host plant (Asclepias curassavica) extract. However, these receptors generally had low sensitivity to three oviposition-stimulating flavonoids identified from this fraction, but they were also sensitive to the butanol fraction of a non-host (Brassica oleracea). Chemoreceptors in sensilla of the tarsomers 2–4 of the mid-legs also responded to the behaviourally active fraction of host plant extract and showed some sensitivity to two of the flavonoids that stimulate oviposition. Similar results were obtained from receptor cells in sensilla on the tip of the antennae. Most of these sensilla had cells responding to the butanol fraction of A. curassavica extract but only 25% of them were also sensitive to one of the behaviourally active flavonoids. These electrophysiological results, in combination with behavioural observations, suggest that host selection in monarch butterflies relies on a complex pattern of peripheral sensory information from several types of tarsal and antennal contact chemoreceptors.     

Developmental Changes of Ecdysteroids in the Eggs of the Silkworm, Bombyx mori

Ecdysteroids were studied in relation to embryonic development and diapause of the silkworm, Bombyx mori . The majority of the ecdysteroids was found to be in the conjugated form, and minor part, in the free form. In the developing eggs, 2-deoxyecdysone, 2-deoxy-20-hydroxy-ecdysone and 3-epi-ecdysone were found to have the free ecdysteroid form as well as the conjugated ecdysteroid form. Ecdysone and 20-hydroxyecdysone almost exclusively had the conjugated form. The concentration of ecdysteroids in the embryo was higher than that in yolkplasm in the early embryonic stages. During the embryonic diapause, the concentration of free ecdysteroids decreased to a low level while the conjugated form maintained the original level.     

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.     

Tritrophic interactions between cabbage cultivars with different resistance and fertilizer levels,cruciferous aphids and parasitoids under field conditions

Tritrophic interactions involving cabbage Brassica oleracea var. capitata cultivars +/- fertilizer, Brevicoryne brassicae (Linnaeus) and Myzus persicae (Sulzer), and the parasitoids Diaeretiella rapae (M'cIntosh) and Aphidius sp. were conducted in 1998 and 1999. Brevicorne brassicae was the dominant aphid species on all cultivars +/- fertilizer, except for some treatments in late season 1998. Ruby Ball (red-leaved with antixenosis factors for B. brassicae alates) +/- fertilizer was consistently less colonized by aphids in early stages of plant growth, although only significantly so compared with Derby Day (green-leaved, susceptible to aphids) without fertilizer for B. brassicae and Minicole (green-leaved with antibiosis factors for B. brassicae) with fertilizer for M. persicae. In early 1999, only B. brassicae was present and no significant differences between cultivars were seen. In the mid to late season 1998, the highest aphid infestations were usually found on Derby Day, although only significantly so for B. brassicae, in some treatments. In 1999, higher aphid infestations were observed on Derby Day in mid to late season and some significant differences were found for M. persicae as well as for B. brassicae. In both years, Ruby Ball had the greatest mummy:aphid ratios early season, with no consistent difference between the other cultivars. Later in the season, mummy:aphid ratios were generally highest on Minicole. Parasitism differed in seasonal occurrence and relative abundance. Diaeretiella rapae mummies were found earlier than Aphidius sp. There was evidence of a beneficial interaction between the degree of plant resistance and biological control in early to mid season.     

Natural Plant Chemicals Acting as Oviposition Deterrents on Cabbage Butterflies (Pieris btassicae (L.), P. rapae (L.) and P. napi (L.))

In several growers' reports Solatium lycopersicum, Sambucus nigra. Thymus vulgaris, Salvia officinalis, Artemisia absinthium, A. abrotanum , and Allium cepa are said to decrease the oviposition of Pieris brassicae, P. rapae and P. napi. In the present study the butterflies were fed with honey automats and reared throughout the year in artificial light in an insec-tarium. In a dual-choice chamber with a slow throughflow of air two equally sized cabbage leaves were placed on opposite sides. Significantly fewer eggs were layed on the cabbage leaf on which extracts of the mentioned plants had been applied. Ten butterflies were used in each experiment. The chemoreceptors and the chemicals involved are not identified but the inhibitory substances are surely secondary plant substances. Acceptance or rejection of secondary plant metabolics determines the complicated food relationships between plants and insects. The use of secondary plant substances for ecological control of insect pests is proposed.     

Circadian orchestration of developmental hormones in the insect, Rhodnius prolixus

This review presents a new perspective on the circadian regulation and functions of insect developmental hormones. In Rhodnius prolixus (Hemiptera), the brain neuropeptide prothoracicotropic hormone (PTTH) is released with a circadian rhythm that is controlled by paired photosensitive clocks in the brain. These clocks comprise the dorsal and lateral PER/TIM clock neurons known to regulate behavioral rhythms in Drosophila. Axons of PTTH and clock cells make close contact. Photosensitive PER/TIM clocks also reside in the paired prothoracic glands (PGs), which generate rhythmic synthesis and release of the ecdysteroid molting hormones. The PG clocks are entrained by both light and PTTH. These four clocks are coupled together by both nerves and hormones into a timing system whose primary regulated output is the circadian rhythm of ecdysteroids in the hemolymph. This complex timing system appears necessary to ensure circadian organization of the gene expression that is induced in target cells by ecdysteroids via circadian cycling of the nuclear ecdysteroid receptor (EcR). This multioscillator system serves to transduce 'the day outside' into endocrine rhythms that orchestrate 'the day inside'. It has many functional similarities with vertebrate circadian systems.     

Position-dependent sensitivity and density of taste receptors on the locust leg underlies behavioural effectiveness of chemosensory stimulation

Chemical stimulation of contact chemoreceptors located on the legs of locusts evokes withdrawal movements of the leg. The likelihood of withdrawal depends on the site of stimulation, in addition to the identity and concentration of the chemical stimulus. A significantly higher percentage of locusts exhibit leg avoidance movements in response to stimulation of distal parts of the leg with any given chemical stimulus compared to proximal sites. Moreover, the percentage of locusts exhibiting avoidance movements is correlated with the density and sensitivity of chemoreceptors on different sites of an individual leg. The effectiveness of chemical stimulation also differs between the fore and hind legs, with NaCl evoking a higher probability of leg withdrawal movements on the foreleg. Moreover, sucrose was less effective than NaCl at evoking withdrawal movements of the foreleg, particularly at low concentrations. The gradients in behavioural responses can be partially attributed to differences in the responsiveness and density of the contact chemoreceptors. These results may reflect the different specialization of individual legs, with the forelegs particularly involved in food selection.     

Steroid control of muscle remodeling during metamorphosis in Manduca sexta

During metamorphosis in the tobacco hornworm, Manduca sexta, the abdominal body-wall muscle DEO1 is remodeled to form the adult muscle DE5. The degeneration of muscle DEO1 involves the dismantling of its contractile apparatus followed by the degeneration of muscle nuclei. As some nuclei are degenerating, others begin to incorporate 5-bromodeoxyuridine (BrdU), indicating the onset of nuclear proliferation. This proliferation is initially most evident at the site where the motoneuron contacts the muscle remnant. The developmental events involved in muscle remodeling are under the control of the steroid hormones, the ecdysteroids. The loss of the contractile elements of the larval muscle requires the rise and fall of the prepupal peak of ecdysteroids, whereas the subsequent loss of muscle nuclei is influenced by the slight rise in ecdysteroids seen after pupal ecdysis. Incorporation of BrdU by muscle nuclei depends on both the adult peak of the ecdysteroids and contact with the motoneuron. Unilateral axotomy blocks proliferation within the rudiment, but it does not block its subsequent differentiation into a very thin muscle in the adult. © 1996 John Wiley & Sons, Inc.     

Ecdysteroids in helminths and annelids

Summary

This review focuses on recent research developments in the past three years concerning ecdysteroid biochemistry in helminths and annelids, with emphasis on results reported at the IXth Ecdysone Workshop in Paris, France, in September 1989. During the past three years, most of the research in this area has been concentrated on nematodes, in several species of which the occurrence of ecdysteroids has been demonstrated. Interesting biological effects of exogenously applied ecdysteroids have been discovered on meiotic reinitiation in oocytes of the dog heartworm, Dirofilaria immitis, and on microfilarial production by Brugia pahangi. These effects, together with previous discoveries, demonstrate the feasibility of affecting nematode physiology with exogenously applied ecdysteroids. However, experiments with four species of nematodes in three separate laboratories have failed to demonstrate that nematodes biosynthesize ecdysteroids from cholesterol. Therefore, it remains to be proven whether ecdysteroids are truly endogenous nematode hormones or are merely compounds with strong biological activity. Research with cestodes and trematodes has similarly revealed that ecdysteroids occur in these organisms and that the compounds have possible regulatory roles, but experiments demonstrating endogenous biosynthesis have yet to be performed. Annelids, which are more closely related to insects than helminths, also contain ecdysteroids; recent research has demonstrated the C-20 hydroxylation of ecdysone to 20-hydroxyecdysone, the active moulting hormone of most insects. Experiments to demonstrate this metabolic step in helminths have failed.