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PubMed1976

Current excitation threshold in sensory neurons of leech central nervous system.

1. Sensory cells in segmental ganglia of the leech (Hirudo medicinalis L.), N (noxious), P (pressure), and T (touch) cells, were stimulated by linearly rising currents. The electrical response to intracellular stimulation with prolonged square-wave currents, the shape of the action potential, and the degree of repetitive activity were also examined. 2. The thresholds for first production of an action potential by linearly rising currents (expressed as multiples of rheobase, I/IO), plotted as a function of the action-potential latency, provide a measure of accomodation. These T-L curves show that the P and T cells accommodate more rapidly than the N cells. 3. An index of accommodation, the accommodation coefficient (I/Io)s/2, was defined as the current required to elicit a first action potential 0.5 s after the beginning of a linearly rising stimulus current. These accommodation coefficients also reflect the property that P and T cells accommodate more rapidly, on the average, than N cells. 4. Threshold depolarization is higher with linearly rising currents than with square-wave currents. 5. Comparison of the accommodation rates of these sensory somata with the published data for other somata and axons, on the basis of accommodation coefficient, minimal current gradient, and/or threshold-latency curve, shows that a) N cells, at the slow end of the leech-soma accommodation range, accommodate at about the same rate as most vertebrate neurons; and b) some vertebrate neurons have accomodation rates as high as or higher than those of the T and P cells. The accommodation rates of most vertebrate neurons are related in a characteristic way to undershoot shape, repetitive-firing properties, and adaptation. Comparable relationships are found in the leech if the N and T cells alone are considered, but the P cells (with extremely rapid accomodation) depart notably from this pattern--they adapt slowly, fire repetitively in response to a maintained stimulus, and have an undershoot of intermediate amplitude and duration.

Adaptation, Physiological↗

PubMed1986

Role of central interneurons in habituation of swimming activity in the medicinal leech.

Swimming activity evoked by light tactile stimulation of a body wall flap in dissected leech preparations undergoes habituation (5). In this study, we examine the activity of several interneurons (cell 204, cell 205, the S cell, and cell 208) during habituation trials to study further the neuronal mechanisms that mediate this decline in responsiveness. Light tactile stimulation of the leech body wall evoked initially a marked excitatory response in cell 204 homologs (segmental swim-initiating neurons) that preceded the initiation of swimming activity. This response decreased over the course of repeated stimulus trials; however, no marked decline in cell 204 activity accompanied the cessation of swim initiation. A similar activity pattern was observed in cell 205. Thus the habituation of swimming activity to stroking of the body wall is not due solely to reduced input to cell 204 and cell 205. The early activity of cell 204 was not correlated to the duration of subsequent swim episodes. However, the impulse frequency of cell 204 during swim episodes was negatively correlated to the period of swim cycles. This correlation between cell 204 activity and cycle period occurred both within individual episodes as well as between trials in a habituation series. Direct stimulation of cell 204 with current pulses evoked swimming activity reliably for an average of 72 trials. Therefore, habituation that results from stroking the body wall (which occurs after approximately 6 trials) is not mediated by plasticity in the connections between cell 204 and the swim oscillator. The S cell fired repeatedly in response to light tactile stimulation. This response declined with repeated trials. Intense intracellular stimulation of the S cell was sufficient to initiate swimming activity in some preparations. The magnitude and duration of the excitation required to initiate swimming by this means were far greater, however, than that which occurred during stroking the body wall. The response of cell 208 (a swim oscillator cell) to body wall stimulation during habituation trials was variable; usually an initial hyperpolarization was followed by some depolarization. No aspect of this response correlated with the onset of habituation. Our results are consistent with the idea that cell 204 and cell 205 are part of the pathway that mediates swimming activity in response to light tactile stimulation of the leech body wall, and that habituation occurs, in part, as the result of reduced sensory input to this cell.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗

PubMed1992

FMRFamide effects on membrane properties of heart cells isolated from the leech, Hirudo medicinalis.

1. The effects of the cardioactive peptide FMRFamide were tested on enzymatically dissociated muscle cells isolated from hearts of the leech. These cells were normally quiescent, with resting potentials near -60 mV. 2. Superfusion of FMRFamide induced a strong depolarization in isolated heart cells (e.g., greater than 40 mV with 10(-6) M FMRFamide). The depolarization was maintained in the continued presence of peptide and persisted long after its removal. Less frequently, FMRFamide superfusion elicited an episodic polarization rhythm. 3. The response of isolated heart cells to bath-applied FMRFamide showed a 1- to 2-min latency. The latency decreased with repeated applications of FMRFamide. 4. The FMRFamide response was diminished by Na+ replacement but persisted with Ca2+ channel blockade. 5. In voltage-clamped heart cells (-60 mv), superfusion of FMRFamide elicited a slow inward current with a transient and a sustained component. 6. Current-voltage (I-V) curves during FMRFamide superfusion in normal leech saline showed that FMRFamide also enhanced voltage-dependent outward currents activated at depolarized levels. 7. Under conditions in which K+ currents were substantially blocked, the FMRFamide-dependent I-V curve was net inward from -90 to +50 mV. A voltage-dependent component was blocked by Co2+ and a linear component by Na+ replacement. 8. We conclude that FMRFamide elicits a persistent inward current with a Na+ component and in addition modulates both voltage-dependent Ca2+ and K+ currents that may contribute to the normal myogenic activity of leech heart muscle cells.

Animals↗

PubMed1999

Modulatory effects of myomodulin on the excitability and membrane currents in Retzius cells of the leech.

Ion channel modulation by the peptide myomodulin (MM) has been demonstrated in a wide variety of organisms including Aplysia, Lymnaea, and Pleurobranchaea. This neural and muscular modulation has been shown to be important for shaping and modifying behavior. In this paper, we report that MM modulates several distinct ionic channels in another species, the medicinal leech Hirudo medicinalis. Experiments have focused on the Retzius cell (R) because the R cell is a multifunction neuron that has been implicated in a number of behaviors including feeding, swimming, secretion, thermal sensing, and the touch elicited shortening reflex and its plasticity. Previous work had identified a MM-like peptide in the leech and demonstrated that this peptide modulated the excitability of the R cell. Using combined current- and voltage-clamp techniques to examine the effects of MM on the R cell, we found that in response to a step pulse, MM increased the excitability of the R cell such that the cell fires more action potentials with a shorter latency to the first action potential. We found that this effect was mediated by the activation of a Na+-mediated inward current near the cell resting membrane potential. Second, we found that MM differentially modulated the potassium currents IA and IK. No effect of MM was found on IA, whereas MM significantly reduced both the peak and steady-state amplitudes of IK by 49 +/- 2.9% and 43 +/- 7.2%, respectively (means +/- SE). Finally we found that MM reduced the amplitude of the Ca2+ current by approximately 20%. The ionic currents modulated by MM are consistent with the overall effect of MM on the cellular activity of the R cell. An understanding of the cellular mechanisms by which MM modulates the activity of the R cell should help us to better understand the roles of both MM and the R cell in a variety of behaviors in the leech.

Animals↗

PubMed2001

Distributed motor pattern underlying whole-body shortening in the medicinal leech.

Whole-body shortening was studied in the leech, Hirudo medicinalis, by a combination of videomicroscopy and multielectrode recordings. Video microscopy was used to monitor the animal behavior and muscle contraction. Eight suction pipettes were used to obtain simultaneous electrical recordings from fine roots emerging from ganglia. This vital escape reaction was rather reproducible. The coefficient of variation of the animal contraction during whole-body shortening was between 0.2 and 0.3. The great majority of all leech longitudinal motoneurons were activated during this escape reaction, in particular motoneurons 3, 4, 5, 8, 107, 108, and L. The firing pattern of all these motoneurons was poorly reproducible from trial to trial, and the coefficient of variation of their firing varied between 0.3 and 1.5 for different motoneurons. The electrical activity of pairs of coactivated motoneurons did not show any sign of correlation over a time window of 100 ms. Only the left and right motoneurons L in the same ganglion had a correlated firing pattern, resulting from their strong electrical coupling. As a consequence of the low correlation between coactivated motoneurons, the global electrical activity during whole-body shortening became reproducible with a coefficient of variation below 0.3 during maximal contraction. These results indicate that whole-body shortening is mediated by the coactivation of a large fraction of all leech motoneurons, i.e., it is a distributed process, and that coactivated motoneurons exhibit a significant statistical independence. Probably due to this statistical independence this vital escape reaction is smooth and reproducible.

Animals↗

PubMed1991

The history of leeching and hirudin.

Leeching is an art dating back at least to ancient Egypt. It reached its zenith in the late 18th and early 19th centuries. The antithrombotic quality of leech saliva was first noted by Haycraft in 1884 and the active anticoagulant ingredient isolated in 1904 by Jacoby. He gave this agent the name 'hirudin'. Hirudin was isolated in pure crystalline form by Markwardt in 1957 and first produced in quantity by genetic engineering in 1986. The salivary glands of several species of leech also contain other biologically active substances which are currently undergoing investigation.

Animals↗

PubMed1992

Use of the medicinal leech in the treatment of ear diseases.

Medicinal leeches (Hirudo medicinalis) and the native secretion of their salivary glands diluted with saline (5 times) were successfully used for the treatment of some ear diseases: tinnitus caused by inner-ear affections, acute external otitis and chronic otitis media. The effect of diluted leech saliva injected into the region of the mastoid process by microelectrophoresis was 25-30% lower than that of the medicinal leeches.

Acute Disease↗

PubMed2004

Evolutionary diversification of specification mechanisms within the O/P equivalence group of the leech genus Helobdella.

Developmental fates and cell lineage patterns are highly conserved in the teloblast lineages that give rise to the segmental ectoderm of clitellate annelids. But previous studies have shown that the pathways involved in specification of the ventrolateral O lineage and the dorsolateral P lineage differ to some degree in distantly related clitellate species such as the leeches Helobdella and Theromyzon, and the sludgeworm Tubifex. To examine this developmental variation at a lower taxonomic level, we have explored the specification pathways of the O and P lineages in the leech genus Helobdella. In leech, the O and P lineages arise from a developmental equivalence group of O/P teloblasts. In this study, we demonstrate that the cell-cell interactions involved in cell fate specification of the O/P equivalence group differ among three laboratory colonies of closely related species. In two populations, the Q lineage is necessary to specify the P fate in the dorsalmost O/P lineage, but in the third population the P fate can be specified by a redundant pathway involving the M lineage. We also observe interspecific variation in the role played by cell interactions within the O/P equivalence group, and in the apparent significance of extrinsic signals from the micromere cell lineages. Our data suggest that cell fate specification in the O/P equivalence group is a complex process that involves multiple cell-cell interactions, and that the developmental architecture of the O/P equivalence group has undergone evolutionary diversification in closely related species, despite maintaining a conserved morphology.

Animals↗

PubMed1993

Lox10, a member of the NK-2 homeobox gene class, is expressed in a segmental pattern in the endoderm and in the cephalic nervous system of the leech Helobdella.

A novel leech homeobox gene, Lox10, is shown to encode a homeodomain sequence characteristic of a phyletically widespread NK-2 homeobox gene class. Lox10 expression was examined in leech embryos of various ages by in situ hybridization. In the unsegmented cephalic region, Lox10 RNA is expressed in a subset of the cells descended from the a' and b' micromeres, including a small cluster of cells, believed to be postmitotic neurons, within the supraesophageal ganglion of the central nervous system. Hybridization signal was not detected in either the mesoderm or ectoderm of the trunk segments, and the apparent restriction of Lox10 ectodermal expression to the nonsegmented cephalic domain resembles the restricted forebrain expression pattern of its mammalian homologues. Lox10 is also expressed within the endodermal tissues of the leech midgut, which arises by cellularization from a polynucleate syncytium. Endodermal expression is organized into a pattern of transverse stripes and spots which are aligned with the intersegmental septa, and which prefigure the pattern of gut wall constrictions observed at later stages of development. Lox10 is the first molecular marker of segmentally periodic endoderm differentiation reported for any animal species.

Amino Acid Sequence↗

PubMed1995

Expression of the cell cycle control gene, cdc25, is constitutive in the segmental founder cells but is cell-cycle-regulated in the micromeres of leech embryos.

The identifiable cells of leech embryos exhibit characteristic differences in the timing of cell division. To elucidate the mechanisms underlying these cell-specific differences in cell cycle timing, the leech cdc25 gene was isolated because Cdc25 phosphatase regulates the asynchronous cell divisions of postblastoderm Drosophila embryos. Examination of the distribution of cdc25 RNA and the zygotic expression of cdc25 in identified cells of leech embryos revealed lineage-dependent mechanisms of regulation. The early blastomeres, macromeres and teloblasts have steady levels of maternal cdc25 RNA throughout their cell cycles. The levels of cdc25 RNA remain constant throughout the cell cycles of the segmental founder cells, but the majority of these transcripts are zygotically produced. Cdc25 RNA levels fluctuate during the cell cycles of the micromeres. The levels peak during early G2, due to a burst of zygotic transcription, and then decline as the cell cycles progress. These data suggest that cells of different lineages employ different strategies of cell cycle control.

Amino Acid Sequence↗

PubMed1997

A nanos homolog in leech.

From the glossiphoniid leech Helobdella robusta, we have cloned and determined the complete coding sequence of Hro-nos, a gene homologous to the nanos gene from Drosophila melanogaster. Developmental northern blots show that Hro-nos, like nanos, is a maternal transcript that decays rapidly during early development. A polyclonal antiserum raised against the HRO-NOS protein was used in developmental western blots and for immunostaining leech embryos of different developmental stages. The HRO-NOS protein is first detectable in 2-cell embryos (4-6 hours of development) and exhibits a transient expression peaking during fourth cleavage (9-12 cells; 8-14 hours of development). The HRO-NOS protein exhibits a graded distribution along the primary embryonic axis and is partitioned unequally between the sister cells DNOPQ and DM, progeny of macromere D' at fourth cleavage: DNOPQ is the segmental ectoderm precursor cell and exhibits levels of HRO-NOS protein that are at least two-fold higher than in cell DM, the segmental mesoderm precursor cell. The observed expression pattern suggests that Hro-nos plays a role in the decision between ectodermal and mesodermal cell fates in leech.

Amino Acid Sequence↗

PubMed2001

Establishment of segment polarity in the ectoderm of the leech Helobdella.

The segmented ectoderm and mesoderm of the leech arise via a stereotyped cell lineage from embryonic stem cells called teloblasts. Each teloblast gives rise to a column of primary blast cell daughters, and the blast cells generate descendant clones that serve as the segmental repeats of their particular teloblast lineage. We have examined the mechanism by which the leech primary blast cell clones acquire segment polarity - i.e. a fixed sequence of positional values ordered along the anteroposterior axis of the segmental repeat. In the O and P teloblast lineages, the earliest divisions of the primary blast cell segregate anterior and posterior cell fates along the anteroposterior axis. Using a laser microbeam, we ablated single cells from both o and p blast cell clones at stages when the clone was two to four cells in length. The developmental fate of the remaining cells was characterized with rhodamine-dextran lineage tracer. Twelve different progeny cells were ablated, and in every case the ablation eliminated the normal descendants of the ablated cell while having little or no detectable effect on the developmental fate of the remaining cells. This included experiments in which we specifically ablated those blast cell progeny that are known to express the engrailed gene, or their lineal precursors. These findings confirm and extend a previous study by showing that the establishment of segment polarity in the leech ectoderm is largely independent of cell interactions conveyed along the anteroposterior axis. Both intercellular signaling and engrailed expression play an important role in the segment polarity specification of the Drosophila embryo, and our findings suggest that there may be little or no conservation of this developmental mechanism between those two organisms.

Animals↗

PubMed1987

Neurite outgrowth and synapse formation by identified leech neurones in culture.

After injury, neurones in the central nervous system (CNS) of the leech regenerate with a high degree of specificity. The aim of our experiments has been to study the sequential steps involved in neurite growth and synapse formation using isolated identified neurones in culture. An important requirement for sprouting of leech neurones is the substrate. Neurites grow only slowly and sparsely on polylysine or vertebrate laminin. The extracellular matrix of leech ganglion capsules contains a protease-sensitive factor which can be extracted with urea. With this material as substrate, growth proceeds rapidly in defined medium. Another neurite-promoting substrate is provided by the plant lectin concanavalin A (Con A). The activity of Con A, but not of the capsule matrix factor, is blocked by the Con A-specific hapten methyl alpha-D-mannoside. The morphology and branching pattern of the neurites in culture depend on the specific substrate and on the type of neurone. During stimulation, less Ca2+ uptake occurs into growth cones than in cell bodies. The mechanism of neurite growth seems not to depend on activity-mediated Ca2+ influx or on interactions between neuronal cell surfaces. However, even without profuse outgrowth, electrical and chemical synapses develop between neighbouring neurones. The type of synapse depends predictably on the types of neurones within the cell pair. Since the development of a synapse can be followed with time in culture, the sequential events can each be studied separately for this multi-step process.

Animals↗

PubMed1988

Excitability and secretory activity in the salivary gland cells of jawed leeches (Hirudinea: Gnathobdellida).

Thousands of salivary cells fill the interstices throughout the anterior ends of jawed leeches. The somata are large (30-200 micron in diameter). They project single processes (ductules) into the three jaws, and were found to fire overshooting action potentials of 50-85 mV amplitude and 100-200 ms duration at low spontaneous rates. The action potentials were not detected in the presence of cobalt (10 mmol l-1), but could be recorded when sodium was absent from the Ringer, so they appear to be calcium-dependent. Salivary material is transported by the long processes of these unicellular glands and secreted into ducts which alternate with paired teeth on the jaws. Secretion is activated reliably by 10(-6) mol l-1 serotonin, but not by other neurotransmitters found in the leech nervous system. Each jaw secretes at an average rate of 230 nl min-1 in the presence of serotonin, and secretion is completely abolished by cobalt. Perfusion with serotonin excites the salivary gland cells into impulse activity, and often evokes bursting. Impulse activity of the peripherally projecting, serotonergic Retzius cells evokes both depolarizations and action potentials in the salivary gland cells. In jawed leeches, central neurones appear to control salivation by a peripheral release of serotonin. This neurotransmitter evokes calcium-dependent action potentials and calcium, in turn, stimulates secretion.

Action Potentials↗

PubMed1997

Chloride secretion drives urine formation in leech nephridia.

The transport mechanisms underlying urine formation in leech nephridia were investigated in situ and in isolated preparations using pharmacological, electrophysiological and micropuncture techniques. Canalicular cells, which secrete the primary urine, function as a Cl(-)-secreting epithelium. An apical Cl- conductance contributes to the lumen-negative potential which drives transcellular K+ transport and paracellular Na+ transport. On the basolateral side, a ouabain-sensitive Na+/K(+)-ATPase contributes substantially to the cellular and transcellular potential and provides the Na+ gradient necessary for a bumetanide-sensitive Na+/K+/2Cl- cotransport. Final urine is formed by subsequent reabsorption of ions along the central canal, where KCl and NaCl are reabsorbed in different portions. The postprandial diuresis is not a consequence of the changes in blood osmolality or ion concentrations. Similar changes in the ionic environment do not promote diuresis in isolated nephridia. Apparently, the composition and volume of the primary urine cannot be separately controlled. Any increase in fluid secretion by leech canalicular cells involves upregulation of the paracellular pathway and stimulation of Cl- entry, which thereby changes the normally K(+)-enriched primary urine to the Na(+)-enriched primary urine characteristic of leeches in diuresis.

Animals↗

PubMed1978

Quantitative mapping of cutaneous receptive fields in normal and operated leeches, Limnobdella.

1. The receptive fields and physiological properties of the sensitive cutaneous mechanoreceptive neurones in the leech Limnobdella australis were found to be very similar to those previously described in Hirudo medicinalis. 2. Following separation from the central nervous system (C.N.S.), the distal dendrite stump and cutaneous receptive field remained unchanged for at least 160 days. 3. There was little spreading of receptive fields into regions of skin isolated from the C.N.S. for at least 184 days. 4. Cutting one dendrite of a mechanoreceptive neurone which has two major dendrites produced little change in the receptive field of the intact dendrite. 5. Abnormalities were found in most of the receptive fields of operated leeches, irrespective of the site of operation. These abnormalities were not seen in normal leeches.

Animals↗

PubMed2005

Morphology and aspects of growth of a trypanosome transmitted by the marine leech Johanssonia arctica (Piscicolidae) from Northern Norway.

The fish leech Johanssonia arctica (Johansson, 1898) was collected from king crabs Paralithodes camtschaticus (Tilesius, 1815) in Finnmark, N Norway, and allowed to feed on experimental fish hosts in the laboratory. This leech ingested blood from laboratory-reared cod (Gadus morhua) and halibut (Hippoglossus hippoglossus). Some experimental halibut acquired trypanosome infection, with parasitaemia between ca. 500 and 60,000 trypanosomes ml(-1). The trypanosomes were of variable size and measured 39-90 microm (mean 57 microm) ca. 81 days post-infection. Characteristic features are cell striation, refractile cytoplasmic granules, anterior nucleus and a relatively long (ca. 6 microm, max 9 microm) distance from the posterior end to the kinetoplast. Following growth, the trypanosomes became increasingly slender, with fewer striae and a nucleus position less pronounced anterior. The trypanosome is considered distinct from a type transmitted by the leech Calliobdella nodulifera (Malm, 1863) in the NE Atlantic, but is regarded conspecific with a trypanosome transmitted by J. arctica in the NW Atlantic. This trypanosome has in the past been identified as Trypanosoma murmanensis Nikitin, 1927, a poorly described species. T. murmanensis cannot be recognized with certainty among the trypanosomes transmitted by C. nodulifera and J. arctica respectively. We propose that the J. arctica-transmitted species is considered T. murmanensis Nikitin, 1927 sensu stricto.

Analysis of Variance↗

PubMed1994

A Kazal-type inhibitor of human mast cell tryptase: isolation from the medical leech Hirudo medicinalis, characterization, and sequence analysis.

Human tryptase, a tetrameric proteinase expressed by mast cells, is virtually unique among the serine proteinases as it is not inhibited by any proteinaceous inhibitor tested so far. We have now isolated, sequenced, and characterized an inhibitor of human tryptase from the medical leech Hirudo medicinalis. LDTI (Leech-Derived Tryptase Inhibitor) was purified to apparent homogeneity by cation exchange and affinity chromatography. Amino acid sequencing of the protein consisting of 46 residues (M(r) 4738) revealed a high degree of similarity to the non-classical Kazal-type inhibitors bdellin B-3 and rhodniin, inhibitors isolated from the medical leech and the insect Rhodnius prolixus, respectively. LDTI is a tight-binding and relatively specific inhibitor of human tryptase; it inhibits only trypsin (EC 3.4.21.4) and chymotrypsin (EC 3.4.21.1) with similar affinities. Inhibition studies using small chromogenic substrates revealed that LDTI inhibits the amidolytic activity of tryptase by approximately 50%, suggesting that most likely due to steric hindrance LDTI binds to and inhibits only 2 of 4 active sites of tryptase. LDTI appears useful as a prototype of inhibitors of human tryptase and as a pharmacological tool for the investigation of the role of tryptase in health and disease.

Amino Acid Sequence↗