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At least 631 records · Page 35Linked to original sources
PubMed1995

Recombinant leech antiplatelet protein specifically blocks platelet deposition on collagen surfaces under flow conditions.

Salivary glands of the leech Haementeria officinalis contain a protein, leech antiplatelet protein (LAPP). This protein was cloned and expressed in yeast and blocks collagen-mediated platelet aggregation and the adhesion of platelets to collagen-coated plates under static conditions. In the current study we investigated the effect of rLAPP on platelet deposition to collagen and collagen-rich surfaces under flow conditions. rLAPP completely inhibited platelet adhesion on collagen types I, III, and IV with IC50 values of 70, 600, and 90 nmol/L, respectively (shear rate = 1600 s-1). Approximately 10-fold more rLAPP was required to obtain a similar inhibition at a low shear rate of 375 s-1. rLAPP caused a concentration-dependent inhibition of binding of 125I-von Willebrand factor (vWF) to collagen type III and was able to displace prebound vWF even after 24 hours. Since platelet adhesion at low shear rate is less dependent on vWF than at high shear rate, this property of rLAPP may explain why less rLAPP is needed at high shear rate than at low shear rate to produce the same effect. Platelet adhesion to collagen type VI was only partially inhibited by rLAPP (maximal 44% with 3 mumol/L rLAPP). rLAPP also caused a pronounced inhibition of platelet deposition to cross sections of human atherosclerotic coronary arteries but had no effect on matrices of cultured human umbilical vein endothelial cells. rLAPP is a potent platelet adhesion inhibitor at high shear rate, which binds to collagen and works by inhibiting binding of vWF to collagen.

Animals↗
PubMed1990

Localization of carbonic anhydrase in identified glial cells of the leech central nervous system: application of histochemical and immunocytochemical methods.

We localized the enzyme carbonic anhydrase (CA) in frozen sections of the leech (Hirudo medicinalis) central nervous system by two histochemical techniques and the indirect immunofluorescence technique. Hansson's cobalt precipitation method and the use of 1-dimethylamino-naphthalene-5-sulfonamide (DNSA) to build a fluorescent enzyme-substrate complex showed that glial cells are the sites of CA activity in the leech. Neuropil and connective glial cells surrounding the axons had strong CA activity, whereas packet glial cells, which surround neuron cell bodies, and neurons themselves remained unstained. Glial cells reacted markedly with FITC-coupled antibodies against CA isoenzyme II, but experiments with antibodies against CA isoenzyme I showed no reaction.

Animals↗
PubMed2004

Cell interactions that affect axonogenesis in the leech Theromyzon rude.

The leech nervous system comprises a relatively simple network of longitudinal (connective) and transverse (segmental) nerves. We have followed the normal pattern of axon development in the glossiphoniid leech Theromyzon rude by immunostaining embryonic preparations with antibody to acetylated alpha-tubulin. The dependence of the normal pattern of axon growth on cells in the mesodermal (M) and ectodermal (N, O, P and Q) lineages was examined by selectively ablating subsets of these lineages in developing embryos. We found that ablating mesoderm severely disrupted overall axonogenesis, while various ectodermal ablations induced a range of more specific phenotypes. In particular, formation of the posterior segmental nerve (PP) was abnormal in embryos deficient in primary neuroectoderm (N lineage). More specific ablations demonstrated that a subset of N-derived cells were required for establishing the PP nerve root. Previous studies have shown that the PP nerve root is normally pioneered by an O lineage-derived neuron (PD). Our results suggest that the role of the N lineage-derived cells is to induce the migration of neuron P(D) to its normal position in the posterior compartment of the hemiganglion.

Animals↗
PubMed2005

Applications of mRNA injections for analyzing cell lineage and asymmetric cell divisions during segmentation in the leech Helobdella robusta.

Synthetic mRNAs can be injected to achieve transient gene expression even for 'non-model' organisms in which genetic approaches are not feasible. Here, we have used this technique to express proteins that can serve as lineage tracers or reporters of cellular events in embryos of the glossiphoniid leech Helobdella robusta (phylum Annelida). As representatives of the proposed super-phylum Lophotrochozoa, glossiphoniid leeches are of interest for developmental and evolutionary comparisons. Their embryos are suitable for microinjection, but no genetic approaches are currently available. We have injected segmentation stem cells (teloblasts) with mRNAs encoding nuclear localized green fluorescent protein (nGFP) and its spectral variants, and have used tandem injections of nGFP mRNA followed by antisense morpholino oligomer (AS MO), to label single blast cell clones. These techniques permit high resolution cell lineage tracing in living embryos. We have applied them to the primary neurogenic (N) lineage, in which alternate segmental founder cells (nf and ns blast cells) contribute distinct sets of progeny to the segmental ganglia. The nf and ns blast cell clones exhibit strikingly different cell division patterns: the increase in cell number within the nf clone is roughly linear, while that in the ns clone is almost exponential. To analyze spindle dynamics in the asymmetric divisions of individual blast cells, we have injected teloblasts with mRNA encoding a tau::GFP fusion protein. Our results show that the asymmetric divisions of n blast cells result from a posterior shift of both the spindle within the cell and the midbody within the mitotic spindle, with differential regulation of these processes between nf and ns.

Animals↗
PubMed1992

Lox2, a putative leech segment identity gene, is expressed in the same segmental domain in different stem cell lineages.

The segmented tissues of the adult leech arise from a set of five, bilaterally paired embryonic stem cells via a stereotyped sequence of cell lineage. Individual segments exhibit unique patterns of cell differentiation, and previous studies have suggested that each stem cell lineage establishes at least some aspects of its own segmental specificity autonomously. In this paper, we describe a putative leech segment identity gene, Lox2, and examine its expression in the various stem cell lineages. Both sequence analysis and the segmental pattern of Lox2 expression suggest a specific homology to the fruitfly segment identity genes Ubx and abdA. In situ hybridization reveals a cellular accumulation of Lox2 RNA over a contiguous domain of 16 midbody segments (M6-M21), including postmitotic neurons, muscles and the differentiating genitalia. Lox2 transcripts were not detected at the stage when segment identities are first established, suggesting that Lox2 gene products may not be part of the initial specification process. Individual stem cell lineages were labeled by intracellular injection of fluorescent tracers, and single cell colocalization of lineage tracer and hybridization reaction product revealed expression of Lox2 RNA in the progeny of four different stem cells. The segmental domain of Lox2 RNA was very similar in the various stem cell lineages, despite the fact that some stem cells generate one founder cell/segment, whereas other stem cells generate two founder cells/segment.

Amino Acid Sequence↗
PubMed1993

Developmental origin of segmental identity in the leech mesoderm.

Segmentation in the leech embryo is established by a stereotyped cell lineage. Each of the 32 segments arises from homologous, bilaterally symmetrical complements of mesodermal and ectodermal blast cell clones. Although segments are homologous, they are regionally differentiated along the longitudinal body axis. Various segments display idiosyncratic ensembles of features, which constitute discrete segmental identities. The differentiation of segment-specific features, such as the mesoderm-derived nephridia, genital primordia and identified Small Cardioactive Peptide immunoreactive neurons, reflects a diversification of the developmental fates of homologous blast cell clones. We have investigated whether segment-specific differentiation of homologous mesodermal blast cell clones depends on cell-intrinsic mechanisms (based on the cells' lineage history) or on cell-extrinsic mechanisms (based on the cells' interactions with their environment) in embryos of Theromyzon rude. For this purpose, we first mapped the segment-specific fates of individual mesodermal blast cell clones, and then induced mesodermal clones to take part in the formation of segments for which they are not normally destined. Two types of ectopic segmental position were produced: one in which a mesodermal blast cell clone was out of register with all other consegmental cells and one in which a mesodermal blast cell clone was out of register with its overlying ectoderm, but was in normal register with the mesoderm and ectoderm on the other side of the embryo. Mesodermal blast cell clones that developed in either type of ectopic segmental position gave rise to segment-specific features characteristic of their original segmental fates rather than their ectopic positions. Thus, the development of segmental identity in the leech mesoderm is attributable to a cell-intrinsic mechanism and, either before or soon after their birth, mesodermal blast cells are autonomously committed to segment-specific fates.

Animals↗
PubMed1994

An axial domain of HOM/Hox gene expression is formed by morphogenetic alignment of independently specified cell lineages in the leech Helobdella.

The homeobox gene Lox2, a member of the HOM/Hox gene class, is expressed in a restricted domain along the anteroposterior (A-P) body axis of the leech Helobdella. The segmental tissues of the leech embryo arise from the parallel merger of five distinct and bilaterally paired cell lineages generated by embryonic stem cells or teloblasts. Injection of cell lineage tracers coupled with anti-LOX2 immunochemistry reveals that all five teloblast lineages generate central nervous system neurons that express the LOX2 protein, and that each lineage expresses LOX2 within a similar domain of body segments. Some lineally identified neurons display anti-LOX2 immunoreactivity over the entire expression domain, but the OM7 neuron has a distinctively high level of LOX2 expression, which is restricted to the seventh midbody ganglion. To ascertain the role of positional information in the axial patterning of LOX2 expression, we performed focal cell ablations that displaced one or another of the teloblast lineages out of segmental register with the other axial tissues. Such displacements brought about a corresponding shift in the LOX2 expression of the perturbed lineage, and had little or no effect on the LOX2 expression of the other, unperturbed lineages. This result indicates that the axial domain of LOX2 expression is not specified by positional cues acting coordinately across the various teloblast lineages, nor would it seem that the expression domain is imprinted from one lineage to the others. Rather, the different teloblast lineages acquire their axial patterns independently, and secondarily bring these patterns into alignment along the A-P axis through a process of morphogenetic assembly.

Animals↗
PubMed1996

Cellular mechanisms of epiboly in leech embryos.

Gastrulation in leech embryos is dominated by the epibolic movements of two tissues: germinal bands, composed of segmental precursor cells, and an overlying epithelium that is part of a provisional integument. During gastrulation, the germinal bands move over the surface of the embryo and coalesce along the prospective ventral midline. Concurrently, the epithelium spreads to cover the embryo. We have begun to analyze the mechanisms involved in gastrulation in the leech by assessing the independent contributions of the epithelium and the germinal bands to these cell movements. Here we describe cellular events during epiboly in normal embryos and in embryos perturbed by either reducing the number of cells in the epithelium, or by preventing the formation of the germinal bands, or both. These experiments indicate that both the germinal bands and the epithelium are able to undergo epibolic movements independently, although each is required for the other to behave as in control embryos.

Animals↗
PubMed2002

Expression and function of an even-skipped homolog in the leech Helobdella robusta.

We have identified homologs of the Drosophila pair-rule gene even-skipped in the glossiphoniid leeches Helobdella robusta and Theromyzon trizonare. In leech embryos, segments arise sequentially from five pairs of embryonic stem cells (teloblasts) that undergo iterated divisions to generate columns (bandlets) of segmental founder cells (primary blast cells), which in turn generate segmentally iterated sets of definitive progeny. In situ hybridization revealed that Hro-eve is expressed in the teloblasts and primary blast cells, and that these transcripts appear to be associated with mitotic chromatin. In more advanced embryos, Hro-eve is expressed in segmentally iterated sets of cells in the ventral nerve cord. Lineage analysis revealed that neurons expressing Hro-eve arise from the N teloblast. To assess the function of Hro-eve, we examined embryos in which selected blastomeres had been injected with antisense Hro-eve morpholino oligonucleotide (AS-Hro-eve MO), concentrating on the primary neurogenic (N teloblast) lineage. Injection of AS-Hro-eve MO perturbed the normal patterns of teloblast and blast cell divisions and disrupted gangliogenesis. These results suggest that Hro-eve is important in regulating early cell divisions through early segmentation, and that it also plays a role in neuronal differentiation.

Amino Acid Sequence↗
PubMed2002

Micromere lineages in the glossiphoniid leech Helobdella.

In leech embryos, segmental mesoderm and ectoderm arise from teloblasts by lineages that are already relatively well characterized. Here, we present data concerning the early divisions and the definitive fate maps of the micromeres, a group of 25 small cells that arise during the modified spiral cleavage in leech (Helobdella robusta) and contribute to most of the nonsegmental tissues of the adult. Three noteworthy results of this work are as follows. (1) The c"' and dm' clones (3d and 3c in traditional nomenclature) give rise to a hitherto undescribed network of fibers that run from one end of the embryo to the other. (2) The clones of micromeres b" and b"' (2b and 3b in traditional nomenclature) die in normal development; the b" clone can be rescued to assume the normal c" fate if micromere c" or its clone are ablated in early development. (3) Two qualitative differences in micromere fates are seen between H. robusta (Sacramento) and another Helobdella sp. (Galt). First, in Helobdella sp. (Galt), the clone of micromere b" does not normally die, and contributes a subset of the cells arising exclusively from c" in H. robusta (Sacramento). Second, in Helobdella sp. (Galt), micromere c"' makes no definitive contribution, whereas micromere dm' gives rise to cells equivalent to those arising from c"' and dm' in H. robusta (Sacramento).

Animals↗
PubMed2004

A cephalic projection neuron involved in locomotion is dye coupled to the dopaminergic neural network in the medicinal leech.

It is widely appreciated that the selection and modulation of locomotor circuits are dependent on the actions of higher-order projection neurons. In the leech, Hirudo medicinalis, locomotion is modulated by a number of cephalic projection neurons that descend from the subesophageal ganglion in the head. Specifically, descending brain interneuron Tr2 functions as a command-like neuron that can terminate or sometimes trigger fictive swimming. In this study, we demonstrate that Tr2 is dye coupled to the dopaminergic neural network distributed in the head brain. These findings represent the first anatomical evidence in support of dopamine (DA) playing a role in the modulation of locomotion in the leech. In addition, we have determined that bath application of DA to the brain and entire nerve cord reliably and rapidly terminates swimming in all preparations exhibiting fictive swimming. By contrast, DA application to nerve cords expressing ongoing fictive crawling does not inhibit this motor rhythm. Furthermore, we show that Tr2 receives rhythmic feedback from the crawl central pattern generator. For example, Tr2 receives inhibitory post-synaptic potentials during the elongation phase of each crawl cycle. When crawling is not expressed, spontaneous inhibitory post-synaptic potentials in Tr2 correlate in time with spontaneous excitatory post-synaptic potentials in the CV motor neuron, a circular muscle excitor that bursts during the elongation phase of crawling. Our data are consistent with the idea that DA biases the nervous system to produce locomotion in the form of crawling.

Action Potentials↗
PubMed1985

Habituation of swimming activity in the medicinal leech.

Tactile stimulation (light stroking) of a body wall flap attached to the ventral nerve cord of the medicinal leech evokes episodes of swimming activity. This swimming response undergoes habituation, involving changes in swim initiation and swim maintenance. Repeated stimulation of the body wall flap evoked swimming activity between three and 39 times before this response failed. During repetitive stimulation, the length of swim episodes decreased by about 50%. The number of swim episodes which could be elicited was not correlated with swim episode length. Following habituation, swim initiation showed significant spontaneous recovery, but swim episode length returned only to 60% of control values. In preparations where spontaneous recovery was followed by rehabituation, the number of swim episodes elicited declined with each habituation-recovery sequence. Additional stimulation immediately following habituation trials had a dual effect: recovery of the swimming response was delayed, but the lengths of swim episodes following spontaneous recovery were increased. Pinching the body wall flap immediately restored the swimming response in an habituated preparation. Swim initiation habituated more rapidly during stimulation of anterior body wall flaps than during stimulation of mid-body or posterior flaps. However, swim length was independent of this regional variation in swim responsiveness. The number of swim episodes elicited by stimulation of body wall flaps attached to posterior or anterior segments depended upon whether this segment was stimulated before or after other flaps. In contrast, in mid-body segments there was no evidence for such stimulus generalization. The lengths of swim episodes elicited during sequential stimulation of several body wall flaps were independent of the stimulation sequence. We propose that separate processes control swim initiation and swim maintenance. These processes must be repeated in most, if not all, of the segmental ganglia of the leech ventral nerve cord.

Action Potentials↗
PubMed1989

Na+, K+ and Ca2+ currents in identified leech neurones in culture.

1. Na+, K+ and Ca2+ currents have been measured by voltage-clamp in Retzius (R), anterior pagoda (AP) and sensory (pressure, touch and nociceptive) cells dissected from the central nervous system (CNS) of the leech. These cells maintain their distinctive membrane properties and action potential configurations in culture. Currents carried by the individual ions were analysed by the use of channel blockers and by their kinetics. Since the cells are isopotential they can be voltage-clamped effectively. 2. Depolarization, as expected, gave rise to an early inward Na+ current followed by a delayed outward K+ current. In Na+-free medium containing tetraethylammonium (TEA+), and in the presence of 4-aminopyridine (4-AP), inward Ca2+ currents were revealed that inactivated slowly and were blocked by Cd2+ and Mn2+. 3. Na+ and Ca2+ currents were similar in their characteristics in R. AP and sensory neurones. In contrast, K+ currents showed marked differences. Three principal K+ currents were identified. These differed in their time courses of activation and inactivation and in their responses to Ca2+ channel blockers. 4. K+ currents of the A-type (IA) activated and inactivated rapidly, were not affected by Ca2+ channel blockers and were eliminated by steady-state inactivation at holding potentials of -30 mV. A-type K+ currents were found in AP cells and as a minor component of the outward current in R cells. A Ca2+-activated K+ current (IC), that inactivated more slowly and was reduced by Ca2+ channel blockers, constituted the major outward current in R cells. The third K+ current resembled the delayed rectifier currents (IK1 and IK2) of squid axons with slow activation and inactivation kinetics. Such currents were found in R cells and in the sensory neurones (T, P and N). 5. The principal differences in membrane properties of identified leech neurones can be explained in terms of the numbers of Na+ channels and the distinctive kinetics of K+ channels in each type of cell.

4-Aminopyridine↗
PubMed1989

Serotonin differentially modulates two K+ currents in the Retzius cell of the leech.

The effects of 100 mumol l-1 serotonin (5-HT) were investigated on the Na+- and Ca2+-dependent action potential and distinct K+ currents in the Retzius (R) cells of the hirudinid leeches Macrobdella decora and Hirudo medicinalis by conventional current-clamp and two-microelectrode voltage-clamp techniques. 1. In normal Na+-containing Ringer, 5-HT decreased the duration of the action potential prolonged by 5 mmol l-1 tetraethylammonium (TEA+) chloride. 2. In Na+-free saline containing 25 mumol l-1 TEA+ to block IK, 5-HT reduced the amplitude and duration of Ca2+ spikes evoked by intracellular current injection. 3. Under voltage-clamp, 5-HT enhanced the peak amplitude of an early transient 4-aminopyridine (4-AP)-sensitive, voltage-dependent outward current, termed IA. A small but significant increase in the time constant of inactivation (tau off) of IA was also measured after exposure to 5-HT. 4. 5-HT suppressed the peak and steady-state amplitudes of a delayed TEA+-sensitive, voltage-dependent outward current, termed IK. These results demonstrate differential simultaneous modulation of distinct K+ currents in the Retzius cell of the leech by the endogenous transmitter serotonin. These cells contain and release 5-HT, and are believed to be multifunction neurons implicated in feeding and swimming. This modulation may change the excitable properties of the cell, leading to a negative feedback autoregulation of its transmitter output.

Action Potentials↗
PubMed1992

Development of segment- and target-related neuronal identity in the medicinal leech.

The rhythmic pumping of the paired heart tubes in the medicinal leech Hirudo medicinalis offers an excellent system for studying the development of a simple behavior in terms of its neuronal and muscular components. The present experiments examined the development of identified heart excitor (HE) motor neurons during normal embryogenesis. Using intracellular impalements and dye-filling, we found that the HE motor neurons could be identified at an early stage of development and that they initially elaborated axonal arborizations in inappropriate target fields in the ventral body wall. These inappropriate projections were retracted as those at the appropriate target (developing heart tube muscle) extended. This remodelling occurred at least 4 days before the HEs acquired the adult phenotype of being driven to fire action potentials in a rhythmic pattern. Although the HEs exhibited centrally driven rhythmic oscillations late in embryogenesis, at earlier stages they exhibited largely a tonic discharge interrupted by bursts of inhibitory potentials in a periodic, but not a rhythmic, pattern. We also found what appeared to be non-rhythmic HE homologs in anterior and posterior segments where HE neurons have not been previously described. These homologs may project along similarly patterned guidance cues early in development, since they are at first indistinguishable from the definitive HEs, but they continued to elaborate both lateral and medial body wall projections over the same period that definitive HEs were expanding their arborizations over the developing heart tube and retracting their body wall projections. In both adult and embryonic leeches the homologs exhibited mostly tonic activity that was interrupted by pronounced, but non-rhythmic, hyperpolarizing postsynaptic potentials. Thus, there appears to be early segmental specification directing the final phenotype of the iterated neuron that, in most segments, becomes the HE motor neuron.

Animals↗
PubMed1993

Segment-specific modulation of the electrophysiological activity of leech Retzius neurons by acetylcholine.

1. The acetylcholine responses of Retzius neurons were electrophysiologically and pharmacologically characterized in situ and in culture. Single-electrode voltage-clamp was used to record currents from leech Retzius neurons from standard segments [Rz(X)] and from reproductive segments [Rz(5,6)]. 2. A 1 s pressure pulse of acetylcholine (ACh) produced a fast inward current followed by a slower outward current in Rz(X) neurons, whereas it produced only an outward current in Rz(5,6) neurons. These segment-specific responses were maintained when the two types of Retzius neurons were isolated in culture for up to 12 days. 3. The inward current of Rz(X) reversed at around -25 mV and was partially carried by Na+. This cationic current desensitized rapidly. The outward current of Rz(X) and Rz(5,6) neurons reversed at around -65 mV and was carried by Cl-. This anionic current desensitized very slowly upon prolonged applications of ACh. 4. The expression of the ACh-induced outward current in Rz(X) was season-dependent and was recorded in a larger proportion of Rz(X) neurons during the summer than during the winter. The expression of the ACh-induced outward current in Rz(5,6) did not show any seasonal pattern. 5. The fast inward current of Rz(X) was also elicited by nicotine; it was blocked by d-tubocurarine, hexamethonium and mecamylamine, but was not affected by alpha-bungarotoxin. The outward current of Rz(X) and Rz(5,6) was also elicited by nicotine and by 4-[N-(3-chlorophenyl)carbamoxyloxy]2-butynyltrimethylammonium chloride (a muscarinic agonist); it was blocked by d-tubocurarine and by alpha-bungarotoxin, but it was not affected by hexamethonium or mecamylamine. 6. The results show that the serotonergic Retzius neurons of the leech could be tonically inhibited by ACh. In addition, the Retzius neurons from standard segments could also be phasically excited by ACh. The receptors responsible for the excitation fit into the classification of neuronal nicotinic receptors, whereas the receptors mediating the inhibition are closer in type to the muscular nicotinic receptor.

(4-(m-Chlorophenylcarbamoyloxy)-2-butynyl)trimethy↗
PubMed1995

Ingestive sensory inputs excite serotonin effector neurones and promote serotonin depletion from the leech central nervous system and periphery.

Thermal and chemical stimuli known to promote ingestive behaviours in the medicinal leech Hirudo medicinalis were tested for their physiological effects on Retzius neurones and for their biochemical effects on serotonin levels in the central nervous system, pharynx and body wall. Retzius neurones throughout the leech nerve cord receive excitatory synaptic input during thermal or chemical stimulation of the prostomial lip. These neurones respond to the rate of change of temperature as well as to absolute temperature at the lip. Exposure of the lip to sodium chloride excites Retzius neurones, whereas exposure to arginine has little effect. Thermal stimulation of the lip elicits a more rapid but less prolonged excitation of Retzius neurones than does chemical stimulation. Stimulation of the prostomial lip is associated with afferent activities in the cephalic nerves D1, D2 and V1-2. Thermal stimulation of the prostomial lip results in depletion of serotonin from midbody ganglia, whereas chemical stimulation has no effect. Conversely, chemical stimulation of the lip results in depletion of serotonin from the body wall, whereas thermal stimulation does not. Pharyngeal serotonin content is decreased with either modality. These data distinguish two important feeding-related sensory input pathways to central serotonergic effector neurones in Hirudo medicinalis.

Animals↗
PubMed1997

Interneuronal and motor patterns during crawling behavior of semi-intact leeches.

Semi-intact tethered preparations were used to characterize neuronal activity patterns in midbody ganglia of the medicinal leech during crawling. Extra- and intracellular recordings were obtained from identified interneurons and from motor neurons of the longitudinal and circular muscles during crawling episodes. Coordinated activities of nine excitatory and inhibitory motor neurons of the longitudinal and circular muscles were recorded during the appropriate phases of crawling. Thus, during crawling, the leech uses motor output components known to contribute to other types of behavior, such as swimming or the shortening/local bending reflex. Interneurons with identified functions in these other types of behavior exhibit membrane potential oscillations that are in phase with the behavior pattern. Therefore, the recruitment of neuronal network elements during several types of behavior occurs not only at the motor neuron level but also involves interneurons. This applies even to some interneurons that were previously thought to have dedicated functions (such as cells 204 and 208 and the S cell). The function of neuronal circuitries in producing different types of behavior with a limited number of neurons is discussed.

Animals↗