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PubMed1990

Seasonal variation of serotonin content and nonassociative learning of swim induction in the leech Hirudo medicinalis.

It is possible to obtain habituation of swim induction by stimulating the leech with repetitive light electrical trains. After obtaining this simple form of non-associative learning, it is also possible to potentiate its response by a series of nociceptive skin brushings (dishabituation). Serotonin applied to the animal is the only neurotransmitter found to mimick dishabituation. We have observed that in the period April-June most animals did not exhibit potentiation of the swimming response after nociceptive stimulation while injection of serotonin mimicked dishabituation as in the animals treated in the period October-March. We have seen correlation between the changes in nonassociative learning and the seasonal variation of serotonin levels in segmental ganglia. This finding strengthens the hypothesis of serotonin as the neurotransmitter mediating dishabituation in swim induction of the leech.

Animals↗
PubMed1993

ATP-inhibited and Ca(2+)-dependent K+ channels in the soma membrane of cultured leech Retzius neurons.

The properties of one ATP-inhibited and one Ca(2+)-dependent K+ channel were investigated by the patch-clamp technique in the soma membrane of leech Retzius neurons in primary culture. Both channels rectify at negative potentials. The ATP-inhibited K+ channel with a mean conductance of 112 pS is reversibly blocked by ATP (Ki = 100 microM), TEA (Ki = 0.8 mM) and 10 mM Ba2+ and irreversibly blocked by 10 nM glibenclamide and 10 microM tolbutamide. It is Ca2+ and voltage independent. Its open state probability (Po) decreases significantly when the pH at the cytoplasmic face of inside-out patches is altered from physiological to acid pH values. The Ca(2+)-dependent K+ channel with a mean conductance of 114 pS shows a bell-shaped Ca2+ dependence of Po with a maximum at pCa 7-8 at the cytoplasmic face of the membrane. The Po is voltage independent at the physiologically relevant V range. Ba2+ (10 MM) reduces the single channel amplitude by around 25% (ATP, TEA, glibenclamide, tolbutamide, and Ba2+ were applied to the cytoplasmic face of the membrane). We conclude that the ATP-dependent K+ channel may play a role in maintaining the membrane potential constant--independently from the energy state of the cell. The Ca(2+)-dependent K+ channel may play a role in generating the resting membrane potential of leech Retzius neurons as it shows maximum activity at the physiological intracellular Ca2+ concentration.

Adenosine Triphosphate↗
PubMed1982

Fluorescent properties of monoamine neurons following glyoxylic acid treatment of intact leech ganglia.

The SPG modification (de la Torre and Surgeon 1976) of the glyoxylic acid method for amine condensation is a straightforward procedure which can be used upon intact ganglia from the leech. Intense fluorescence of the neurosomata of identified neurons which contain either indoleamine (serotonin, 5-HT) or catecholamine (CA) is obtained in less than 30 min. The fluorescence of the 5-HT containing neurons is yellow (518-526 nm) and decays more rapidly than the dominant blue emission (478-480 nm) of the CA neurons. Most importantly, the SPG technique greatly enhances the visibility of the axonal processes of neurons which contain both classes of amines over the fluorescence produced by formaldehyde condensation techniques. Both blue and yellow fluorescent varicosities are easily distinguished in the longitudinal connectives and lateral roots of the leech C.N.S. Because of its simplicity and high fluorescence yields, the SPG method for histochemical fluorescence should contribute to investigations of amine functions in invertebrate nervous systems.

Animals↗
PubMed1996

CCD imaging of the electrical activity in the leech nervous system.

A single ganglion of the nervous system of the leech Hirudo medicinalis was isolated. One or both roots emerging from each side of the ganglion were sucked into suction pipettes used either for extracellular stimulation or for recording the gross electrical activity. The ganglion was stained with the fluorescence voltage sensitive dye Di-4-Anepps. The fluorescence was measured with a nitrogen cooled CCD camera. Our recording system allowed us to measure in real time slow optical signals corresponding to changes in light intensity of at least 5/1000. These signals were caused by the direct polarization of neuronal structures, the afterhyperpolarization or the afterdischarge induced by a prolonged stimulation. When images were acquired at fixed times, several of them could be averaged and optical signals of at least 2/1000 could be reliably measured. These optical signals originated from well identified neurons, such as T, P and N sensory neurons. By taking images at different times and at different focal planes, electrical events could be followed at a temporal resolution of 50 Hz. The three dimensional dynamics of electrical events, initiated by a specific stimulation, was imaged and the spread of excitation among leech neurons was followed. When two roots were selectively stimulated, their neuronal interactions could be imaged and the linear and non-linear terms of the interaction could be characterized.

Animals↗
PubMed1986

A kinematic study of crawling behavior in the leech, Hirudo medicinalis.

The medicinal leech crawls along a solid substrate by repeated alternating extensions and shortenings of the body. Extension occurs with the posterior sucker attached and the head sucker free. The head sucker then attaches, followed by shortening and release of the tail sucker. The tail sucker is then pulled toward the head, where it reattaches to the substrate. The head sucker then releases, and another crawling cycle begins (Figs. 1, 5). There are two crawling variants: inchworm crawling, in which the head and tail suckers are closely apposed at the end of a cycle and the body forms a loop above the substrate, and vermiform crawling, in which the suckers are placed farther apart and the body remains fairly close to the substrate (Fig. 1). The cycle period and the distance traveled during a cycle are greater in inchworm than in vermiform crawling; however, the velocity of travel is the same for both (Fig. 2). For both variants, the interval between head sucker attachment and tail sucker release is similar at all cycle periods and has a value consistent with direct interneuronal conduction of a signal from head sucker sensory neurons to tail sucker motor neurons. The interval between tail sucker attachment and head sucker release, however, is longer and varies with the cycle period, suggesting a more complex interneuronal circuit in the pathway from tail sucker sensory neurons to head sucker motor neurons (Fig. 4). The onsets of the components of the crawling cycle (extension, post-extension pause, shortening, and post-shortening pause) show an anteroposterior lag (Figs. 5, 7). For both variants, the travel time between segments varies directly with the period (Fig. 8). For both crawl types, the durations of the cycle components vary directly with the period, with several exceptions (Figs. 9, 10). A model is presented that summarizes the coordination of the various motor events in a cycle of leech crawling (Figs. 11 and 12).

Animals↗
PubMed1986

Initiation of swimming activity by trigger neurons in the leech subesophageal ganglion. III. Sensory inputs to Tr1 and Tr2.

In papers I and II of this series, we described two pairs of interneurons, Tr1 and Tr2, in the leech subesophageal ganglion which can trigger swimming activity in the isolated central nervous system (CNS). In this paper, we describe sensory inputs to these trigger neurons from previously identified mechanosensory neurons. We found that: Weak mechanical stimulation (stroking) of a body wall flap attached to a segmental ganglion in an otherwise isolated CNS excites the contralateral Tr1 slightly. Strong mechanical stimulation (pinching) of a mid-body wall flap evokes a burst of impulses in the contralateral Tr1. For both means of stimulation the effects on the ipsilateral Tr1 and on the Tr2 cell pair were much weaker. Stroking a body wall flap attached to the head ganglion (supra- and subesophageal ganglia) in an otherwise isolated CNS excites both Tr1s and both Tr2s, although the effect is weaker for the Tr2s. Pinching strongly excites both trigger neurons bilaterally. Pressure and nociceptive mechanosensory neurons (P and N cells) in the subesophageal ganglion and the first segmental ganglion appear to make direct excitatory synapses with the contralateral Tr1 and Tr2. Mechanosensory interactions with the ipsilateral trigger neurons appear to be indirect. Functional inactivation of Tr1 by hyperpolarization does not prevent swim initiation either by weak mechanical stimulation of a body-wall flap or by intracellular stimulation of P cells.2+ We conclude that the trigger neurons, Tr1 and Tr2, provide an excitatory pathway by which mechanosensory stimulation can initiate leech swimming activity.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗
PubMed1988

Release of octopamine by Leydig cells in the central nervous system of the leech Macrobdella decora, and its possible neurohormonal role.

1. The release of octopamine by the central nervous system of the leech Macrobdella decora was examined. Isolated ganglia or chains of ganglia were incubated in salines of varying composition, and the release of octopamine into the perfusate was measured using a radioenzymatic method. In some experiments this release was correlated with the electrical activity of the octopamine-containing Leydig cells, as measured via a microelectrode in the cell body. 2. Chains of ganglia incubated in normal saline released 0.04 pmol octopamine/ganglion/3 min incubation period. This amount was not significantly increased by either the monoamine oxidase inhibitor iproniazid phosphate (0.1 mM) or the uptake inhibitor desipramine (10 microM) alone, but was by both together. Nominally calcium-free saline containing 20 mM Mg++ significantly decreased octopamine release. 3. High K+ saline increased octopamine release significantly in both standard saline and one containing the blocking agents. This increase was sevenfold in saline containing iproniazid phosphate and desipramine, and was significantly greater than that obtained in saline without the blockers. This provides further evidence for the role of octopamine as a neuroactive substance in the leech by indicating the existence of possible mechanisms for its uptake and/or inactivation. 4. Octopamine release was positively correlated with the firing frequency of Leydig cells. No release was detectable when the cells were prevented from firing by the injection of hyperpolarizing current. Release was frequency-dependent when the cells fired at frequencies of 0.1-1.0 spikes/s. Elevating the external calcium concentration from 1.8 to 5.4 mM significantly increased octopamine release at all frequencies tested, except for 0 spikes/s, at which release remained below detectabilty.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗
PubMed1985

Properties of action potentials carried by divalent cations in identified leech neurons.

Properties of divalent cation potentials carried by either Sr2+ or Ca2+ ions in Na+-free, TEA-Ringer solution were characterized in identified neurons of two species of leeches (Macrobdella and Haementeria). In Macrobdella, the overshoot of the potentials varied logarithmically with [Sr2+]0 (28.5 mV per 10-fold change). The overshoot, Vmax, and duration of the potentials increased with increasing divalent cation concentration and saturated at about 20 to 30 mM [Sr2+]0. The Vmax, amplitude, and duration of the potentials were reversibly blocked by Co2+ and Mn2+. The block by Mn2+ could be well-fitted by a reverse Langmuir-curve with an apparent KI of 100 micromolar. The local anesthetic procaine also reversibly inhibited the Vmax and duration of the potentials. The inhibition was greater at alkaline pH suggesting that procaine blocks the calcium channel from inside the membrane. The identified leech neurons examined in Macrobdella varied considerably in their ability to sustain somatic divalent cation potentials. Stimulation of T cells and most motoneurons produced no or only weak potentials, whereas stimulation of Retzius, N, Nut, and AP cells evoked overshooting potentials of several seconds' duration. Stimulation of the ALG cell of Haementeria in normal Ringer solution evoked a slowly-rising, purely Ca2+-dependent potential of approximately 100 ms duration. This response was TTX-resistant, unaffected by complete removal of Na+ from the Ringer solution, and abolished by 1 mM Mn2+. The overshoot varied logarithmically with a slope of 28 mV/decade change in [Ca2+]0.

Action Potentials↗
PubMed1982

Immunohistochemical demonstration of peptides, serotonin and dopamine-beta-hydroxylase-like material in the nervous system of the leech Hirudo medicinalis.

The central ganglia of the leech, Hirudo medicinalis, were processed for the immunohistochemical localisation of bombesin-, substance P-, cholecystokinin-, vasoactive intestinal polypeptide-, enkephalin-, serotonin- and dopamine-beta-hydroxylase-related substances. To varying extents all of the substances were localised in neuropile processes, and all, with the exception of substance P, were associated with specific perikarya. The most prominent neuropeptides, in terms of the number of immunoreactive neurones, were cholecystokinin and vasoactive intestinal peptide. The dopamine-beta-hydroxylase positive neurones are thought to be octopaminergic, and the serotonin monoclonal antibody revealed positive staining in the Retzius cells. We were unable to demonstrate the coexistence of pairs of substances in any neurones in the leech ganglia.

Animals↗
PubMed1983

Distinctions between gap junctions and sites of intermediate filament attachment in the leech C.N.S.

Freeze-fracture studies on the nerve cord of the leech Hirudo medicinalis reveal that the plasma membranes of various cells, including glial and muscle cells, contain at least two distinct types of aggregated intramembrane particles, identified as hemidesmosomes and gap junctions. Hemidesmosomes consist of angular particles irregularly arranged in circular or elongate patches in external leaflets (E-faces), and are associated with a bundle of intermediate filaments extending into the cytoplasm. Hemidesmosomes of specific axons abut on extracellular space at openings in the surrounding glial sheath. Gap junctions are patches of rounder particles in cytoplasmic leaflets (P-faces) and are more uniformly spaced; they have a corresponding array of pits in the complementary E-face. Gap junctions connect processes of adjacent smooth muscle cells, and apparently interconnect glial processes. Thus, different types of cells in the leech C.N.S. have similar intramembrane specializations. Moreover, the hemidesmosomes and gap junctions might, on superficial examination, be confused.

Animals↗
PubMed1996

Glutamate as a transmitter in the sensory pathway from prostomial lip to serotonergic Retzius neurons in the medicinal leech Hirudo.

The involvement of glutamate in putative ingestive sensory pathways affecting the excitability of serotonergic Retzius neurons (RZ) in the leech CNS was investigated with a pharmacological approach. Exposure of the prostomial lip to 150 mM NaCl and 1 mM arginine produced excitatory as well as inhibitory responses in RZ found in the reproductive segments, while only excitatory responses were elicted in standard midbody RZ. Antagonists of glutamatergic receptors of the kainate/quisqualate type effectively inhibited chemosensory dependent excitation of RZ. Antagonists of glutamatergic receptors of the N-methyl D-aspartate type were ineffective in this regard. Cephalic nerve stimulation, like chemical stimulation of the lip, produced segment-specific responses in midbody RZ. Both the polysynaptic and monosynaptic components of the excitatory response of standard midbody RZ following cephalic nerve stimulation were inhibited in the presence of the kainate/quisqualate antagonist DNQX. These data suggest a role for glutamate as a transmitter in the neural circuitry from receptors of the leech prostomial lip to serotonergic RZ.

Animals↗
PubMed1996

Salivary lipid profiles of the leech (Hirudo medicinalis).

Saliva was collected from sixteen leeches (Hirudo medicinalis). The saliva was analyzed for its total lipid content (3.26 +/- 0.31 mg of total lipids per 100 mL saliva). The lipids were separated into polar and nonpolar by chromatographic techniques. The neutral fraction was approximately 2/3 of the total, and the polar fraction was approximately 1/3 of the total lipids. Thin-layer chromatography was used to obtain the individual profiles of the polar and nonpolar lipids. Of the identified lipids, phosphatidic acids and free fatty acids represented the largest percentage. These results suggest that the leech contains a unique lipid distribution, and that some of these components may be potent phospholipases and lipases that probably are present in its saliva for the purpose of preventing plugging or healing of the wound in the attacked organism.

Animals↗
PubMed2001

Modulation of Ca2+ influx in leech Retzius neurons. I. Effect of extracellular pH.

We investigated the cytosolic free Ca2+ concentration ([Ca2+]i) of leech Retzius neurons in situ while varying the extracellular and intracellular pH as well as the extracellular ionic strength. Changing these parameters had no significant effect on [Ca2+]i when the membrane potential of the cells was close to its resting value. However, when the cells were depolarized by raising the extracellular K+ concentration or by applying the glutamatergic agonist kainate, extracellular pH and ionic strength markedly affected [Ca2+]i, whereas intracellular pH changes appeared to have virtually no effect. An extracellular acidification decreased [Ca2+]i, while alkalinization or reduction of the ionic strength increased it. Correspondingly, [Ca2+]i also increased when the kainate-induced extracellular acidification was reduced by raising the pH-buffering capacity. At low extracellular pH, the membrane potential to which the cells must be depolarized to evoke a detectable [Ca2+]i increase was shifted to more positive values, and it moved to more negative values at high pH. We conclude that in leech Retzius neurons extracellular pH, but not intracellular pH, affects [Ca2+]i by modulating Ca2+ influx through voltage-dependent Ca2+ channels. The results suggest that this modulation is mediated primarily by shifts in the surface potential at the extracellular side of the plasma membrane.

Animals↗
PubMed2003

To swim or not to swim: regional effects of serotonin, octopamine and amine mixtures in the medicinal leech.

Focally treating the head brain of the medicinal leech Hirudo medicinalis with various biogenic amines affected the initiation, termination and maintenance of fictive swimming (i.e., the neural correlate of swimming). Application of serotonin to saline surrounding only the head brain inhibited fictive swimming, whereas removing serotonin induced swimming. This contrasts sharply with previous observations that serotonin applied to the nerve cord induces swimming. Although application of octopamine to the brain activated swimming, a mixture of octopamine and serotonin inhibited swimming. Subsequent removal of this mixture from the brain activated robust swimming and was more potent for activating swimming than either the removal of serotonin or the application of octopamine. Swim episodes induced by brain-specific manipulations of octopamine had more swim bursts per episode than those induced by serotonin. These brain-specific effects of the amines on fictive swimming are probably due to the modulation of higher-order circuits that control locomotion in the leech. We observed that serotonin or a mixture of serotonin and octopamine hyperpolarized an identified descending brain interneuron known as Tr2. Removal of the mixture caused Tr2 to exhibit membrane potential depolarizations that correlated in time with the expression of swim episodes.

Amines↗
PubMedDate not supplied

A central pattern generator underlies crawling in the medicinal leech.

Crawling in the medicinal leech has previously been thought to require sensory feedback because the intact behavior is strongly modulated by sensory feedback and because semi-intact preparations will only crawl if they can move freely. Here we show that an isolated leech nerve cord can produce a crawling motor pattern similar to the one seen in semi-intact preparations, which consists of an anterior-to-posterior wave of alternating excitatory circular and longitudinal motor neuron bursts in each segment. The isolated cord also reproduces the patterns of activity seen in semi-intact preparations for several other kinds of cells: the dorsal inhibitor cell 1, the ventral excitor cell 4, and the annulus erector motor neuron. Because this correspondence is so strong, there must be a central pattern generator in the isolated cord that can produce the basic motor pattern for crawling without sensory feedback. A quantitative analysis of the isolated motor pattern, however, reveals that isolated and semi-intact preparations have longer periods than the intact behavior and that there are deficiencies in the timing of motor neuron bursts in the isolated pattern. These results suggest that sensory feedback modulates the isolated central pattern generator to help produce the normal motor pattern.

Algorithms↗
PubMed2000

Functionally heterogeneous segmental oscillators generate swimming in the medical leech.

Swimming behavior in the leech Hirudo medicinalis arises from neuronal circuits within the ventral nerve cord. Although the ventral nerve cord comprises a series of homologous segmental ganglia, it remains unresolved whether the swim oscillator circuits within individual ganglia are functionally equivalent. We have extended previous studies on pairs of ganglia to test whether individual ganglia throughout the nerve cord are capable of generating swim oscillations and to measure the cycle periods of local oscillations. We found that the swim-generating function of individual ganglia is broadly distributed, but not uniform. The swim-like oscillations in isolated ganglia from the anterior ganglia nerve cord were less robust than those from mid-cord. Swimming activity in posterior cord ganglia is even weaker we were unable to obtain swim-like oscillations from individual ganglia of the nerve cord posterior to segment 12. Swim-cycle periods exhibited a U-shaped function: those recorded in the most anterior individual ganglia (2.3 s for ganglion M2) and short chains of posterior ganglia (up to 4.0 s) were two to four times longer than those obtained from mid-cord ganglia (near 1.0 s). We conclude that the leech swim system comprises a functionally heterogeneous set of local oscillator units.

Animals↗
PubMedDate not supplied

The phosphoinositide signaling cascade is involved in photoreception in the leech Hirudo medicinalis.

The effects of BAPTA, heparin, and neomycin on electrical light responses were studied in the photoreceptors of Hirudo medicinalis. Light activation produces a fast increase in intracellular Ca2+ concentration (Cai) as detected with the fluorescent Ca2+ indicator calcium green-5N. Chelating intracellular calcium by injections of 10 mmol(-1) BAPTA suppresses spontaneous quantum bumps, reduces light sensitivity by more than 2 log(10) units, and substantially increases the latent period of light responses. BAPTA strongly inhibits the plateau phase of responses to long steps of light. Injections of 45-100 mg ml(-1) of heparin act in a similar manner to BAPTA, affecting the latency of the light responses even more. De-N-sulfated heparin, an inactive analog, is almost ineffective at the same concentration compared with heparin. Heparin diminishes the light-induced Cai elevation significantly, whereas de-N-sulfated heparin does not. Intracellular injections of 50-100 mmol l(-1) of the aminoglycoside neomycin, which inhibits phospholipase-C-mediated inositol 1,4,5-trisphosphate formation, acts similar to BAPTA and heparin. Pressure injections of the hydrolysis resistant analog of inositol 1,4,5-trisphosphate, inositol 2,4,5-trisphosphate, strongly depolarize leech photoreceptors and mimic an effect of light adaptation. These results suggest a close similarity between phototransduction mechanisms in leech photoreceptors and existing models for visual transduction in other invertebrate microvillar photoreceptors.

Animals↗
PubMed2001

Leeches in the larynx.

This is a report of four patients who inhaled leeches. They presented with severe attacks of inspiratory stridor, difficulty in breathing and spitting blood. Indirect laryngoscopy revealed a dark greenish living foreign body in the larynx in each case. Laryngoscopy was performed with local anaesthesia in two of the cases and general anaesthesia was used in the other two. The leeches were removed with forceps.

Adult↗