Biomedical subjects

B Leitch

Publications and source records attributed to B Leitch.

At least 19 recordsLinked to original sources

PubMed2000

Snowdrop lectin (GNA) has no acute toxic effects on a beneficial insect predator, the 2-spot ladybird (Adalia bipunctata L.).

Two-spot ladybird (Adalia bipunctata L.) larvae were fed on aphids (Myzus persicae (Sulz.)) which had been loaded with snowdrop lectin (Galanthus nivalis agglutinin; GNA) by feeding on artificial diet containing the protein. Treatment with GNA significantly decreased the growth of aphids. No acute toxicity of GNA-containing aphids towards the ladybird larvae was observed, although there were small effects on development. When fed a fixed number of aphids, larvae exposed to GNA spent longer in the 4th instar, taking 6 extra days to reach pupation; however, retardation of development was not observed in ladybird larvae fed equal weights of aphids. Ladybird larvae fed GNA-containing aphids were found to be 8-15% smaller than controls, but ate a significantly greater number of aphids (approx. 40% to pupation). GNA was shown to be present on the microvilli of the midgut brush border membrane and within gut epithelial cells in ladybird larvae fed on GNA-dosed aphids, although disruption of the brush border was not observed. It is hypothesised that GNA does not have significant direct toxic or adverse effects on developing ladybird larvae, but that the effects observed may be due to the fact that the aphids fed on GNA are compromised and are thus a suboptimal food.

Journal Article↗

PubMed1999

Modulation of transmitter release from the locust forewing stretch receptor neuron by GABAergic interneurons activated via muscarinic receptors.

The role of muscarinic receptors in the down-regulation of acetylcholine (ACh) release from the locust forewing stretch receptor neuron (fSR) terminals has been investigated. Electrical stimulation of the fSR evokes monosynaptic excitatory postsynaptic potentials (EPSPs) in the first basalar motoneuron (BA1), produced mainly by the activation of postsynaptic nicotinic cholinergic receptors. The general muscarinic antagonists scopolamine (10(-6) M) and atropine (10(-8) to 10(-6) M) caused a reversible increase in the amplitude of electrically evoked EPSPs. However, scopolamine (10(-6) M) caused a slight depression in the amplitude of responses to ACh pressure-applied to the soma of BA1. These observations indicate that the EPSP amplitude enhancement is due to the blockade of muscarinic receptors on neurons presynaptic to BA1. The muscarinic receptors may be located on the fSR itself and act as autoreceptors, and/or they may be located on GABAergic interneurons which inhibit ACh release from the fSR. Electron microscopical immunocytochemistry has revealed that GABA-immunoreactive neurons make presynaptic inputs to the fSR. The GABA antagonist picrotoxin (10(-6) M) caused a reversible increase in the EPSP amplitude, which does not appear to be due to an increase in sensitivity of BA1 to ACh, as picrotoxin (10(-6) M) slightly decreased ACh responses recorded from BA1. Application of scopolamine (10(-6) M) to a preparation preincubated with picrotoxin did not cause the EPSP amplitude enhancement normally seen in control experiments; in fact, it caused a slight depression. This indicates that at least some of the presynaptic muscarinic receptors are located on GABAergic interneurons that modulate transmission at the fSR/BA1 synapse.

Acetylcholine↗

PubMed1999

GABA immunoreactivity in processes presynaptic to the locust wing stretch receptor neuron.

Studies on the locust Locusta migratoria (Leitch et al. [1998] Soc. Neurosci. Abstr. 624.10) suggest that gamma-aminobutyric acid (GABA)ergic interneurons, activated via muscarinic receptors, may be involved in modulation of transmitter release at synapses between the forewing stretch receptor (fSR) and wing depressor motoneurons. To help elucidate the role GABAergic interneurons may play in modulation of transmitter release from fSR terminals, the proportion and distribution of GABA-immunoreactive (GABA-IR) inputs to the fSR were analysed using double-labelling (horseradish peroxidase and GABA immunocytochemistry) and electron microscopy. Forty-three percent of synaptic inputs to the fSR were from GABA-IR profiles, the majority of which were located on lateral branches. The highest proportion (57%) of inputs to the fSR, however, were from non-IR processes containing either clear spherical vesicles or mixed clear and dense-cored vesicles. Outputs from the fSR to GABA-IR profiles were also found, although their number was relatively low (7%). Networks were identified in which both the fSR and its non-IR postsynaptic partner received input from the same GABA-IR neuron. Such an arrangement would allow both pre- and postsynaptic inhibition of fSR afferent outputs, for example at fSR/motoneuron synapses. These observations demonstrate that the fSR does receive presynaptic inputs from GABA-IR neurons, thus providing morphological support for pharmacological and electrophysiological findings that GABAergic neurons are involved in the presynaptic modulation of the fSR. Nevertheless, modulation at this synapse may be more intricate and involve other, as yet unidentified, neurotransmitters released from non GABA-IR presynaptic processes and also muscarinic receptors located on the fSR itself.

Animals↗

PubMed1996

GABAergic synapses in the antennal lobe and mushroom body of the locust olfactory system.

To help elucidate the role of inhibitory feedback in the genesis of odour-evoked synchronization of neural activity, we investigated the distribution of gamma-aminobutyric acid (GABA)ergic synaptic terminals in the antennal lobes (AL) and mushroom bodies (MB) of the locust olfactory system. Electron-microscopy, intracellular horseradish peroxidase labelling, and immunocytochemistry were combined to assess the distribution of GABAergic synapses, using established methods (Leitch and Laurent [1993] J. Comp. Neurol. 337:461-470). In the AL, GABA-immunoreactive presynaptic terminals contacted both immunoreactive and immunonegative profiles. Conversely, GABA-immunoreactive profiles received direct input from both reactive and negative terminals. The tract containing the axons of the projection neurons that run from the AL to the MB contained about 830 axons of fairly uniform size, none of which was immunoreactive for GABA. In the calyx of the MB, large immunoreactive terminals contacted very-small-diameter profiles thought to belong to the Kenyon cells (KCs). This was confirmed by combining immunocytochemistry with intracellular HRP-labelling of KCs. KCs were not immunoreactive for GABA. Although some GABAergic contacts were made onto the spiny profiles of KCs, others were made onto their dendritic shafts. Large GABA-immunoreactive profiles were also found to contact large negative profiles that were presynaptic to KC terminals. This suggests that KC dendrites can be both pre- and post-synaptically inhibited in the calyx. The MB pedunculus contained ca. 50,000 tightly packed KC axons, showing conspicuous en passant and often reciprocal synaptic contacts between neighbouring axons. KC axons were immunonegative, but received direct input from, and contacted directly, large immunoreactive profiles running across or along the KC axons. In the alpha- and beta-lobes of the MB, connections similar to those in the pedunculus were seen with two main differences: (1) The density of synaptic profiles was higher, giving on occasion numerous serially connected profiles in a single section; (2) large immunonegative profiles with dense-core vesicles were abundant and were frequently presynaptic to GABAergic processes and to very-small-diameter profiles which possibly belong to KCs. These results are discussed in the context of the known physiological data on olfactory processing in these complex circuits.

Animals↗

PubMed1995

Localization of a nicotinic acetylcholine receptor-like antigen in the thoracic nervous system of embryonic locusts, Schistocerca gregaria.

Sodium dodecyl sulfate-polyacrylamide gel electrophoresis and subsequent immunoblotting of neuronal membrane proteins derived from thoracic ganglia of adult Locusta and Schistocerca reveal that a polyclonal antiserum raised against the Locusta nicotinic acetylcholine receptor (nAChR), binds strongly to an identical polypeptide band corresponding to 65 kDa in both locust species. This polyclonal antiserum was used to analyze the distribution of antigenic sites within the developing thoracic central nervous system of Schistocerca embryos. Axonal outgrowths from the earliest differentiated neurons are first labeled between 30% and 35% development. By 40% to 45% development, labeled granules appear in the cytoplasm of neuronal cell bodies. When the developing neuropil is first enclosed at approximately 45% to 50% development, it appears uniformly labeled, but by 55% development, unlabeled areas appear that represent the sites of future tracts and commissures. By 75%, an adult pattern of neuropil immunogenicity is established in which synaptic regions are stained but tracts and commissures are not. This suggests that during the early development of the thoracic nervous system nAChR-like antigenic sites are evenly distributed, but later become concentrated in the developing synaptic areas.

Animals↗

PubMed1995

Modulation of transmitter release from the terminals of the locust wing stretch receptor neuron by muscarinic antagonists.

The forewing stretch receptor (SR) neuron makes monosynaptic connections with wing depressor motoneurons; in this article the pharmacology of its output onto the first basalar motoneuron (BA1) has been investigated. The SR, like other insect afferents that have been studied so far, appears to be cholinergic; transmission was suppressed reversibly by the nicotinic antagonist gallamine (10(-4) M) and irreversibly by alpha-bungarotoxin (10(-6) M). The choline reuptake blocker hemicholinium-3 (10(-4) M) also caused a reversible reduction in the amplitude of SR excitatory postsynaptic potentials (EPSPs) recorded in BA1. The receptor subtype nonselective muscarinic antagonists atropine (10(-4) M), scopolamine (10(-4) M), and quinuclidinyl benzilate (10(-5) M), unlike nicotinic antagonists, caused an augmentation in EPSP amplitude. This effect does not appear to be caused by an increase in sensitivity of the motoneuron to acetylcholine (ACh), since atropine produced a marked reduction rather than an increase in the amplitude of responses to ACh pressure applied to the soma of BA1. Scopolamine only caused a modest reduction in the amplitude of ACh somatic responses. The simplest explanation for these observations is that muscarinic antagonists bring about an increase in EPSP amplitude by blockade of presynaptic autoreceptors that normally down-regulate the release of ACh from SR terminals. The effects of muscarinic receptor subtype-selective antagonists indicate that presynaptic receptors in this preparation may have a pharmacological profile more similar to that of vertebrate M2 receptors than to that of M1 or M3 subtypes. The functional significance of autoreceptors in this preparation are discussed.

Acetylcholine↗

PubMed1993

Distribution of GABAergic synaptic terminals on the dendrites of locust spiking local interneurones.

Double-labelling and electron microscopy were used to assess the distribution of GABAergic synapses made onto the neurites of spiking local interneurones in the locust. The aims were to determine the sites of inputs mediating inhibition of the spiking local interneurones and to ascertain the relative abundance of such inputs. This information should allow us to understand better the integrative properties of these spiking local interneurones and the role of inhibition in shaping their receptive field properties or in fine tuning their spike-mediated outputs. Spiking interneurones in a midline population were labelled by intracellular injection of horseradish peroxidase after physiological characterisation. Colloidal gold immunocytochemistry was then used on ultrathin sections of these neurones with a polyclonal antibody raised against GABA. Most GABAergic (inhibitory) input synapses onto the interneurones are made on their ventral neurites, which also receive afferent (excitatory) inputs. These inhibitory inputs to the ventral neurites constitute 43% of the identifiable synapses. Relatively few GABAergic inputs were found onto the dorsal neurites, which are predominantly the sites of output synapses from these interneurones. These results suggest that much synaptic integration takes place in the ventral field of branches and that GABA-mediated presynaptic inhibitory control of spike-mediated outputs from the dorsal neurites is unlikely to occur.

3,3'-Diaminobenzidine↗

PubMed1993

Distribution of acetylcholine receptors in the central nervous system of adult locusts.

A polyclonal antibody raised against nicotinic acetylcholine receptor protein from purified locust neuronal membrane was used to analyse the distribution of antigenic sites within the central nervous system of adult Schistocerca gregaria. Light microscopic examination showed that all principal neuropiles in the thoracic ganglia label with the antibody but that the major tracts and commissures do not. Analysis of this pattern of staining in the electron microscope reveals that the receptor is present on specific synaptic and extrajunctional neuronal membranes in the neuropile. Antigenic sites are also evident on the plasma membranes and within the cytoplasm adjacent to Golgi complexes of some neuronal somata, suggesting that these neurones synthesise nicotinic acetylcholine receptors. In addition to neuronal labelling, there is evidence that the receptor is also present on the membranes of three types of glial cells. The implications of this pattern of receptor distribution are discussed.

Animals↗

PubMed1993

GABA-immunoreactivity in processes presynaptic to the terminals of afferents from a locust leg proprioceptor.

Individually labelled sensory neurons from the femoral chordotonal organ, a proprioceptor at the femoro-tibial joint of a locust hindleg, were analysed by intracellular recording, and by electron microscopical immunocytochemistry to reveal the arrangement of their input and output synapses and to determine whether the input synapses were GABAergic. Intracellular recordings from these sensory neurons show spikes superimposed on a barrage of synaptic potentials during movements of the femoro-tibial joint. These synaptic inputs can be mimicked by GABA. Input synapses are made onto the vesicle-containing terminals of afferents and are often closely associated with the output synapses. By contrast, the axons of the afferents in the neuropil have no vesicles and neither make nor receive synapses. The input synapses to the afferent terminals are made from processes typically a few microns in diameter, whereas the output synapses are made onto much smaller processes of only 0.1-0.2 micron. Input synapses at which an afferent terminal is the only postsynaptic element are common. Where the synapse is dyadic the second postsynaptic element does not usually appear to be a chordotonal afferent. The output synapses from the afferent terminals are usually dyadic. At 78% of the input synapses, the presynaptic neurite showed immunoreactivity to a GABA antibody, supporting the physiological evidence that the presynaptic effects can be mediated by the release of GABA. The remaining (22%) immunonegative synapses are intermingled with those showing GABA immunoreactivity, but their putative transmitter is unknown. These morphological observations suggest that the presynaptic control of the chordotonal afferents is largely mediated by GABAergic neurons, but because other types of neuron also appear to be involved, presynaptic modulation may be more complex than has yet been revealed by the physiology.

Action Potentials↗

PubMed1992

Embryonic development of synapses on spiking local interneurones in locust.

The development of synapses on an identified population of spiking local interneurones in the thoracic ganglia of embryonic locusts was examined by means of intracellular horseradish peroxidase injection and electron microscopy. In adult locusts, spiking local interneurones of the midline group receive direct inputs onto a ventral field of branches from leg mechanosensory afferents and in turn make output synapses, mainly from a dorsal field of branches, directly upon leg motor neurones, nonspiking local interneurones, and intersegmental interneurones. The aim of this study is to examine the development of these connections. These interneurones are born relatively late in embryogenesis and are not identifiable until approximately 55% of development. At this time (55-60%) only simple filopodial contacts or punctate contacts are evident between the stained interneurones and other neurones. By 65-70% embryogenesis, vesicles are found adjacent to regions where apposed membranes are symmetrically thickened with amorphous electron-dense material. These symmetrical contacts lack distinct presynaptic bar-shaped densities and therefore, are not considered to be synapses. At this stage, the interneurones do not produce action potentials upon intracellular injection of depolarising current. Morphologically identifiable synapses, with vesicles, a presynaptic bar, and relatively little postsynaptic density, are first evident at 70-75%, coincident with the time of arrival of the majority of leg mechanosensory afferents into the central nervous system. At this stage, action potentials and synaptic potentials are also recorded for the first time. The midline spiking interneurones thus become electrically excitable when synapses are first recognisable, at approximately 70% embryogenesis. Most of the synapses found on the interneurones are outputs. The ratio of outputs to inputs on ventral branches is 7.5:1 which contrasts markedly to the adult ratio of 1:2. By 85-90%, output synapses still predominate on the ventral branches, but the ratio of outputs to inputs is reduced to almost 2:1. Dorsal branches have predominantly output synapses throughout embryogenesis. The ratio of dorsal outputs to inputs at 85-90% is 8.5:1 which compares with the adult ratio of 6.5:1. At this stage, action potentials and synaptic activity are always recorded.

Action Potentials↗

PubMed1992

Two classes of vesicles are present and change in relative proportion during post-embryonic development of rectifying electrical synapses in the crayfish.

The size and shape of vesicles at junctional appositions of the rectifying electrical synapses between the medial giant fibre and motor giant neurone of the crayfish were measured during the first 2 months after hatching. Summed data over this period reveal a bimodal distribution in vesicle diameter. From the day of hatching until about 7 days of age, small vesicles (circa 25 nm diameter) predominate. From day 7 onwards, larger vesicles (circa 55 nm diameter) occur in increasing numbers, until at day 56 they constitute about 85% of the population at any one junctional apposition. At intermediate ages (day 7-28) individual junctional appositions may show the same bimodal distribution in size as does the age group as a whole, indicating that large and small vesicles occur together at the same junction. The larger vesicles are mainly circular, while the small vesicles are pleomorphic, with shapes ranging from almost circular down to a shape factor of about 0.6.

Aging↗

PubMed1992

Structural and functional post-embryonic development of a non-rectifying electrical synapse in the crayfish.

The post-embryonic development of the non-rectifying septate synapse between homologous lateral giant (LG) fibre segments has been investigated using electron microscopy and electrophysiology. In adults, the LG-LG synapse is characterized by closely apposed membranes (approximately 4 nm separation) traversed by regularly spaced particles, and large (60-80 nm) spherical vesicles on both sides of the junction. In newly hatched crayfish the junction between lateral giant fibre segments comprises regions of close membrane apposition as seen in the adult along with non-specialized areas of wide (10-15 nm) membrane separation. Vesicles associated with these junctions are small (25-40 nm) and pleomorphic. The number of vesicles is low by comparison with adult junctions; in most sections of hatchling junctions there are normally fewer than five vesicles, although as many as 30 have occasionally been seen. During development the non-specialized areas of wide membrane separation become rare and the vesicle population changes to a mixture of small pleomorphic forms and larger (60-80 nm) spherical ones. However even at two months the number of large spherical vesicles is markedly less than that at the adult synapse, while small pleomorphic vesicles are still abundant. Despite the difference between the adult and hatchling vesicle populations, intracellular recordings have shown that the synapse is fully functional as a non-rectifying electrical junction on hatching and that the intracellular marker Lucifer Yellow can pass between adjacent lateral giant fibre neurons.

Animals↗

PubMed1992

Ultrastructure of electrical synapses: review.

This article reviews studies providing information on the ultrastructure of electrical synapses. Although the review focuses on electron-microscopic investigations, its aim is to examine how the structure of an electrical synapse relates to its function. It begins by presenting a historical overview of the early studies which were responsible for the recognition of electrical synapses. The structure of gap junctions which are the morphological correlates of electrical synapses is illustrated and the ultrastructure and function of the two types of electrical synapse, rectifying and non-rectifying, described. Recent papers investigating the ultrastructure of electrical and mixed electrical-chemical synapses in invertebrates and vertebrates are reviewed. For earlier references, the reader is directed to previous reviews on the subject. Much new information, however, on the structure and formation of electrical synapses has been obtained from work on cultured neurons and from electron-microscopic, immunocytochemical, conformational and molecular studies. This article reviews those studies and in light of their findings, re-examines the relationships of the structure of electrical synapses with their function.

Animals↗

PubMed1991

Postembryonic development of rectifying electrical synapses in crayfish: physiology.

In a previous paper we showed that the ultrastructure of the giant fibre to motor giant synapse of crayfish changes in the first few weeks after hatching from having predominantly the appearance of a chemical synapse to having the appearance of an electrical synapse. This is paralleled by a behavioural change from non-giant fibre-mediated to giant fibre-mediated tailflips. In this paper we describe the physiology of the giant fibre to motor giant synapse over this period. We find the following: (1) The giant fibre to motor giant synapse usually transmits spikes 1:1 from the day of hatching. (2) The synapse operates by electrical transmission from the day of hatching, when no connexons are apparent at the ultrastructural level. (3) The synapse has no detectable chemical component, even at an age when the predominant type of junctional apposition has the ultrastructural appearance of a chemical synapse. (4) Inhibitory chemical synapses occur onto the motor giant at the day of hatching, and these show similar physiological characteristics to those which occur onto the motor giant in adults. (5) In some preparations, the giant fibre to motor giant electrical synapse shows rectification similar to that in the adult, but in most cases both depolarizing and hyperpolarizing current injected into the medial giant spreads to the motor giant. (6) Current spread from the medial giant to the motor giant is increased by hyperpolarizing the motor giant neuron, even when medial giant to motor giant transmission is apparently non-rectifying. (7) Both the giant fibre and the motor giant have resting potentials of about -90 mV. There is no standing difference in resting potential as there is in the adult. This may explain the apparent lack of medial giant to motor giant rectification observed in most preparations.

Animals↗

PubMed1990

Anti-GABA antibodies label a subpopulation of chemical synapses which modulate an electrical synapse in crayfish.

Antibodies raised against gamma-aminobutyric acid (GABA) were used to stain sections from the crayfish abdominal nervous system, and the sections were examined under the electron microscope using a protein-A/gold conjugate secondary label. Sections were taken through the third ganglionic root, and through the interganglionic connective at the base of the third root posterior to the ganglia. The third root contains two very large motor axons, a non-GABAergic excitor (Motor Giant; MoG), and a GABAergic inhibitor (Flexor Inhibitor; FI). Only one of the two large axons stained positively for GABA, confirming that the antibody has high specificity for GABAergic neurones. The MoG is driven by powerful electrical synapses from the giant fibres, but also receives inhibitory chemical synaptic input which can gate the excitatory input. There is no physiological evidence for any other form of chemical input. However, at the ultrastructural level, the MoG is postsynaptic to three types of chemical profiles; SE-type containing round agranular vesicles, SI-type containing pleomorphic vesicles, and SM-type containing a mixture of round agranular and dense-cored vesicles. There is a highly differentiated staining pattern of these three synaptic types. Only the SI-type profiles stain positively with the GABA antibody, while the SE- and SM-type do not show significant staining. This suggests that the MoG can under some circumstances receive chemical input other than GABAergic inhibitory input. These other types of input have yet to be physiologically identified.

Animals↗

PubMed1990

The gap junction-like form of a vacuolar proton channel component appears not to be an artifact of isolation: an immunocytochemical localization study.

Gap junctional structures containing a 16-kDa intrinsic membrane protein have been isolated from the hepatopancreas of the crustacean Nephrops norvegicus. These structures are double membranes 14-15 nm thick and composed of hexagonal arrays of particles which have a central pore that is penetrated by a cationic negative stain. Membrane preparations have also been isolated from the hepatopancreas and these contain similar gap junctional regions of uniform width. Affinity purified antibodies to the 16-kDa protein bind principally to these gap junctional regions. Antiserum raised against the isolated gap junctional structures binds strongly to the lateral surfaces of the columnar epithelial cells and in particular to gap junction-like regions.

Animals↗

PubMed1990

Schistosoma haematobium: the effect of Astiban on the cell composition and ultrastructure of the vitelline gland and the ultrastructure of the tegument and gastrodermis.

Treatment of Schistosoma haematobium (Nigerian strain) in hamsters with a single dose of 40 mg/kg of Astiban caused a reduction in the number of S1, S2, and S3 vitelline cells and an increase in S4 cells. Following seven daily doses of the drug, a marked reduction in S1 cells and a complete loss of S2 and S3 cells occurred such that 95% of the cells were S4 cells, all of which were structurally abnormal. Coagulation and disintegration of the protein granules of the vitelline droplets occurred with increase in lipid droplets, swelling of the nuclear membrane and an increase in cytosegresomes. Blebbing of the tegument in both sexes occurred following a single treatment and vacuolation of the basal infolds and alterations to the mitochondria also resulted, but severe erosion of the tegument was rare even following repeated drug treatment. Damage to the gastrodermis was severe with the development of autophagic vacuoles containing whorls of myelin and sequestered portions of damaged tissue. The degree of damage increased with the number of drug treatments.

Animals↗

PubMed1989

Post-embryonic development of rectifying electrical synapses in the crayfish: ultrastructure.

The post-embryonic development of the rectifying Giant Fibre-Motor Giant (GF-MoG) synapse and the Giant Fibre-Segmental Giant (GF-SG) synapse has been investigated using electron-microscopy. In adults, the MoG and SG neurons make contact with the GFs by sending narrow 'finger-like' processes through the glial and connective tissue sheath surrounding each GF. The junctional region is characterized by closely apposed membranes (approximately 4 nm separation) traversed by regularly spaced connections, and large (60-80 nm) spherical vesicles in the presynaptic fibre. In newly hatched crayfish junctional contact is made over extensive areas of flat membrane apposition, due to the absence of a thick connective sheath around the giant fibres. Initially the junctional region is dominated by contacts which are morphologically indistinguishable from chemical synapses, i.e. 1. The apposed membranes are separated by a cleft of approximately 20-30 nm (an order of magnitude larger than the cleft distance at electrotonic synapses). 2. There is pre- and post-synaptic thickening of the junctional membranes with a dense cytoplasmic material. 3. Small (25-40 nm) pleomorphic vesicles are found on the presynaptic side of the junction, commonly in association with a dense presynaptic bar. Regions of junctional contact displaying the adult electronic-type morphology first appear at approximately one week post-hatching. At this age they are limited in distribution and occupy a central position in the area of contact surrounded by a broad 'chemical-like' annulus. During subsequent development these sites with electrotonic-type morphology grow in relative size, so that the 'chemical-like' sites become compressed towards the edges of the regions of contact. The adult type of morphology, in which the 'chemical-like' regions are vestigial, is achieved approximately two months after hatching.

Animals↗