Properties of the nociceptive neurons of the leech segmental ganglion

1996 ◽  
Vol 75 (6) ◽  
pp. 2268-2279 ◽  
Author(s):  
J. Pastor ◽  
B. Soria ◽  
C. Belmonte
Keyword(s):  
Receptive Fields ◽  
Noxious Stimulation ◽  
Impulse Frequency ◽  
Time Intervals ◽  
Noxious Heat ◽  
Nociceptive Neurons ◽  
Slow Rise ◽  
Segmental Ganglion ◽  
Heat Sensitization ◽  
Sustained Discharge

1. The electrical responses of nociceptive (N) lateral and N medial neurons of the leech segmental ganglion to mechanical, chemical, and thermal stimulation of the skin were studied in a superfused ganglion-body wall preparation. 2. Mechanical indentation of the skin > 10 mN evoked in both types of cells a sustained discharge of impulses; afterdischarge was often observed with suprathreshold stimulations. 3. Application to the cutaneous receptive area of 10-100 mM acetic acid or of NaCI crystals and solutions also elicited a firing response in N medial and N lateral cells. In contrast, capsaicin applied to the skin (3.3 x 10(-5) to 3.3 x 10(-2) M) excited N lateral but not N medial neurons. Likewise, impulse discharges were obtained when capsaicin was applied to the cell bodies of N lateral but not of N medial neurons. 4. In both types of N neurons, heating of the skin above 39 degrees C evoked a discharge of impulses whose frequency was roughly proportional to temperature values. 5. Application of repeated suprathreshold heating cycles at 10-min intervals enhanced the impulse frequency of the response (sensitization). Shorter time intervals between heating cycles depressed the response to heat. Sensitization could not be obtained by equivalent soma depolarizations obtained by intracellular current injection. 6. Impulse discharges evoked by irritant agents were also augmented by previous application of noxious heat. 7. N lateral neurons fired in response to low-pH solutions and capsaicin directly applied onto the ganglion. N medial neurons responded inconsistently to acid and were insensitive to capsaicin. Action potentials evoked in N lateral cells by capsaicin had a slow rise, a prominent hump, and a prolonged afterhyperpolarization. 8. It is concluded that N neurons of the leech segmental ganglion respond to different modalities of noxious stimuli applied to their peripheral receptive fields and develop sensitization after repeated noxious stimulation. These properties are typical of mammalian polymodal nociceptors; thus N neurons may be a simple model for analysis of membrane mechanisms associated with polymodality of nociceptive neurons.

Responses of primate SI cortical neurons to noxious stimuli

1983 ◽  
Vol 50 (6) ◽  
pp. 1479-1496 ◽  
Author(s):  
D. R. Kenshalo ◽  
O. Isensee
Keyword(s):  
Receptive Field ◽  
Cortical Neurons ◽  
Receptive Fields ◽  
The Body ◽  
Thermal Stimulation ◽  
Discharge Frequency ◽  
Mechanical Stimuli ◽  
Noxious Heat ◽  
Nociceptive Neurons ◽  
Heat Pulses

Recordings were made from single SI cortical neurons in the anesthetized macaque monkey. Each isolated cortical neuron was tested for responses to a standard series of mechanical stimuli. The stimuli included brushing the skin, pressure, and pinch. The majority of cortical neurons responded with the greatest discharge frequency to brushing the receptive field, but neurons were found in areas 3b and 1 that responded maximally to pinching the receptive field. A total of 68 cortical nociceptive neurons were examined in 10 animals. Cortical neurons that responded maximally to pinching the skin were also tested for responses to graded noxious heat pulses (from 35 to 43, 45, 47, and 50 degrees C). If the neuron failed to respond or only responded to 50 degrees C, the receptive field was also heated to temperatures of 53 and 55 degrees C. Fifty-six of the total population of nociceptive neurons were tested for responses to the complete series of noxious heat pulses: 46 (80%) exhibited a progressive increase in the discharge frequency as a function of stimulus intensity, and the spontaneous activity of two (4%) was inhibited. One population of cortical nociceptive neurons possessed restricted, contralateral receptive fields. These cells encoded the intensity of noxious mechanical and thermal stimulation. Sensitization of primary afferent nociceptors was reflected in the responses of SI cortical nociceptive neurons when the ascending series of noxious thermal stimulation was repeated. The population of cortical nociceptive neurons with restricted receptive fields exhibited no adaptation in the response during noxious heat pulses of 47 and 50 degrees C. At higher temperatures the response often continued to increase during the stimulus. The other population of cortical nociceptive neurons was found to have restricted, low-threshold receptive fields on the contralateral hindlimb and, in addition, could be activated only by intense pinching or noxious thermal stimuli delivered on any portion of the body. The stimulus-response functions obtained from noxious thermal stimulation of the contralateral hindlimb were not different from cortical nociceptive neurons with small receptive fields. However, nociceptive neurons with large receptive fields exhibited a consistent adaptation during a noxious heat pulse of 47 and 50 degrees C. Based on the response characteristics of these two populations of cortical nociceptive neurons, we conclude that neurons with small receptive fields possess the ability to provide information about the localization, the intensity, and the temporal attributes of a noxious stimulus.4+.

Responses of neurons in primate ventral posterior lateral nucleus to noxious stimuli

1980 ◽  
Vol 43 (6) ◽  
pp. 1594-1614 ◽  
Author(s):  
D. R. Kenshalo ◽  
G. J. Giesler ◽  
R. B. Leonard ◽  
W. D. Willis
Keyword(s):  
Receptive Fields ◽  
Dorsal Column ◽  
Sensory Cortex ◽  
Noxious Stimulation ◽  
Mechanical Stimuli ◽  
Thalamic Neuron ◽  
Noxious Heat ◽  
Thalamic Neurons ◽  
Noxious Stimuli ◽  
Stimulation Of

1. Recordings were made from the caudal part of the ventral posterior lateral (VPLc) nucleus of the thalamus in anesthetized macaque monkeys. In additon to many neurons that responded only to weak mechanical stimuli, scattered neurons were found that responded to both innocuous and noxious stimulation or just to noxious stimulation of the skin. A total of 73 such neurons were examined in 26 animals. 2. Noxious stimuli included strong mechanical stimuli (pressure, pinch, and squeezing with forceps) and graded noxious heat (from 35 degrees C adapting temperature to 43, 45, 47, and 50 degrees C). The responses of the VPLc neurons increased progressively with greater intensities of noxious stimulation. The stimulus-response function when noxious heat stimuli were used was a power function with an exponent greater than one. 3. Repetition of the noxious heat stimuli revealed sensitization of the responses of the thalamic neurons to such stimuli. The threshold for a response to noxious heat was lowered, and the responses to supra-threshold noxious heat stimuli were enhanced. 4. The responses of VPLc neurons to noxious heat stimuli adapted after reaching a peak discharge frequency. The rate of adaptation was slower for a stimulus of 50 degrees C than for one of 47 degrees C. 5. For the six neurons tested, responses to noxious heat were dependent on pathways ascending in the ventral part of the lateral funiculus contralateral to the receptive field (ipsilateral to the thalamic neuron). In two cases, the input to the thalamic neurons from axons of the dorsal column was also conveyed by way of a crossed pathway in the opposite ventral quadrant. In another case, access to the thalamic neuron by way of ascending dorsal column fibers was demonstrated. 6. The thalamic neurons had restricted contralateral receptive fields that were somatotopically organized. Neurons with receptive fields on the hindlimb were in the lateral part of the VPLc nucleus, whereas neurons with receptive fields on the forelimb were in medial VPLc. 7. Ninety percent of the VPLc neurons tested that responded to noxious stimuli could be activated antidromically by stimulation of the surface of SI sensory cortex. It was possible to confirm that many of these cells project to the SI sensory cortex by using microstimulation. Successful microstimulation points were either within the SI cortex or in the white matter just beneath the cortex. 8. We conclude that some neurons in the VPLc nucleus are capable of signaling noiceptive stimuli. The nociceptive information appears to reach these cells through the ventral part of the lateral funiculus on the side contralateral to the receptive field, presumably by way of the spinothalamic tract. The VPLc cells are somatotopically organized, and they are thalamocortical neurons that project to the VPLc nucleus and SI cortex play a role in nociception.

Role of [Ca2+]i in the ATP-Induced Heat Sensitization Process of Rat Nociceptive Neurons

1999 ◽  
Vol 81 (6) ◽  
pp. 2612-2619 ◽  
Author(s):  
M. Kress ◽  
S. Guenther
Keyword(s):  
Calcium Ionophore ◽  
Noxious Heat ◽  
Nociceptive Neurons ◽  
Adult Rat ◽  
Caged Calcium ◽  
Sensitization Process ◽  
Heat Sensitization ◽  
Inflamed Tissue

Role of [Ca2+]i in the ATP-induced heat sensitization process of rat nociceptive neurons. In inflamed tissue, nociceptors show increased sensitivity to noxious heat, which may account for heat hyperalgesia. In unmyelinated nociceptive afferents in rat skin in vitro, a drop of heat threshold and an increase in heat responses were induced by experimental elevation of intracellular calcium ([Ca2+]i) levels with the calcium ionophore ionomycin (10 μM). Similar results were obtained in experiments employing [Ca2+]irelease from preloaded “caged calcium” (NITR-5/AM) via UV photolysis. In both cases, sensitization was prevented by preventing rises in [Ca2+]i with the membrane-permeant calcium chelator BAPTA-AM (1 mM). No pronounced change of mechanical sensitivity was observed. Heat-induced membrane currents ( I heat) were investigated with patch-clamp recordings, and simultaneous calcium measurements were performed in small sensory neurons isolated from adult rat dorsal root ganglia (DRG). Ionomycin-induced rises in [Ca2+]iresulted in reversible sensitization of I heat. In the same subset of DRG neurons, the endogenous algogen ATP (100 μM) was used to elevate [Ca2+]i, which again resulted in significant sensitization of I heat. In correlative recordings from the skin–nerve preparation, ATP induced heat sensitization of nociceptors, which again could be blocked by preincubation with BAPTA-AM. Rises in [Ca2+]iin response to inflammatory mediators, e.g., ATP, thus appear to play a central role in plastic changes of nociceptors, which may account for hypersensitivity of inflamed tissue.

Neurons in ventrobasal region of cat thalamus selectively responsive to noxious mechanical stimulation

1983 ◽  
Vol 49 (3) ◽  
pp. 662-673 ◽  
Author(s):  
C. N. Honda ◽  
S. Mense ◽  
E. R. Perl
Keyword(s):  
Sciatic Nerve ◽  
Mechanical Stimulation ◽  
Receptive Fields ◽  
High Threshold ◽  
Noxious Stimulation ◽  
Nociceptive Neurons ◽  
Subcutaneous Tissues ◽  
Alpha Beta ◽  
Lateral Nucleus ◽  
Stimulation Of

1. A survey was made of neurons located in the ventral posterior lateral nucleus of the cat thalamus and its immediate vicinity for elements with specifically nociceptive properties. 2. Pipette microelectrodes filled with a dye solution were used to obtain extracellular recordings of unitary activity in 34 animals anesthetized with chloralose. 3. The great majority of the over 1,000 different single units responding to sciatic nerve stimulation noted in this series of experiments could also be excited by innocuous mechanical stimulation of skin or subcutaneous tissues. An infrequent but consistently noted group of units excited by A-alpha beta delta sciatic nerve volleys did not respond to innocuous mechanical manipulation or A-alpha beta sciatic nerve volleys; they were excited only by either noxious levels of mechanical stimulation or when volleys included the activity of more slowly conducting myelinated fibers. The latencies of such "high-threshold" units to sciatic volleys were longer than those of the other units. 4. Histologically identified recording sites marked by dye were recovered for 17 high-threshold units. Twelve of the 17 could be excited by noxious manipulations of restricted parts of the contralateral hindlimb. Nine of the 12 had cutaneous receptive fields, whereas 3 responded only to stimulation of subcutaneous tissues. None of the 17 high-threshold units evidenced additional discharges that could be correlated with the C-fiber component of sciatic nerve volleys. 5. The high-threshold units typically exhibited a low level of irregular background activity, which increased on repeated noxious stimulation of the peripheral receptive fields. Tactile units of the same or adjacent penetrations usually had a much greater degree of ongoing activity, often marked by bursts at a relatively high frequency. 6. The recording sites for the 17 high-threshold neurons were located dorsal and ventrolateral to the core of the ventrobasal nuclei and were not found in the midst of the low-threshold, cutaneous, mechanoreceptive population. During vertical stereotaxic penetrations, high-threshold units were noted dorsal or ventral to the location of ventrobasal tactile units in a pattern consistent with the core's somatotopic arrangement. 7. These results support the concept that the cat ventrolateral thalamus receives a small but distinct selectively nociceptive projection. The nociceptive neurons appear to be located in a shell that surrounds the main tactile projection to the ventral posterior lateral nucleus and that retains at least part of the topographic arrangement characteristic of the tactile core. Presumably, this projection is part of an organization identifying and localizing noxious stimulation.

Responses of neurons in the lateral cervical nucleus of the cat to noxious cutaneous stimulation

1987 ◽  
Vol 57 (6) ◽  
pp. 1686-1704 ◽  
Author(s):  
K. C. Kajander ◽  
G. J. Giesler
Keyword(s):  
Receptive Fields ◽  
Noxious Stimulation ◽  
Mechanical Stimuli ◽  
Cutaneous Stimulation ◽  
Nociceptive Neurons ◽  
Nociceptive Responses ◽  
Lateral Cervical Nucleus ◽  
Noxious Mechanical Stimulation ◽  
Thermal Stimuli ◽  
Stimulation Of

The majority of neurons at the origin of the spinocervical tract are driven by noxious stimulation of their receptive fields. Surprisingly, previous studies have encountered only a small percentage of nociceptive neurons within the terminus of the spinocervical tract, the lateral cervical nucleus (LCN). To determine if previous reports have underestimated the proportion of nociceptive LCN neurons, 129 neurons within the nucleus were physiologically identified and examined in cats prepared using three different methods. Fifty-nine percent of the neurons studied in unanesthetized cats that were decerebrated and spinalized responded either differentially or exclusively to noxious mechanical stimulation of the skin within discrete receptive fields. LCN neurons also gave accelerating responses to increasingly more intense noxious thermal stimuli. LCN neurons are, therefore, capable of coding both the intensity and location of noxious stimuli. Only 6% of LCN neurons responded to noxious cutaneous stimuli in unanesthetized, decerebrated cats in which the spinal cord was intact. Only 4% of LCN neurons in intact urethan-anesthetized cats were driven by noxious stimulation. Several previous studies of the LCN have been performed in cats that were deeply anesthetized with barbiturates. Therefore, the effects of barbiturates on the nociceptive responses of LCN neurons were determined. Subanesthetic doses of intravenously administered barbiturates reduced or eliminated the responses of nociceptive LCN neurons to noxious thermal stimuli in decerebrated and spinalized cats. Responses to innocuous mechanical stimuli by these neurons were not blocked by barbiturates. Nociceptive LCN neurons in decerebrated and spinalized cats were somatotopically organized. Neurons with forelimb receptive fields were located in the ventromedial half of the LCN; neurons with hindlimb receptive fields were located in the dorsolateral half of the nucleus. This report and previous studies of the spinocervical tract suggest that the spinocervicothalamic pathway is capable of playing an important role in nociception.

Nociceptive neurons in area 24 of rabbit cingulate cortex

1992 ◽  
Vol 68 (5) ◽  
pp. 1720-1732 ◽  
Author(s):  
R. W. Sikes ◽  
B. A. Vogt
Keyword(s):  
Body Surface ◽  
Response Latency ◽  
Receptive Fields ◽  
Dorsal Surface ◽  
Response Duration ◽  
Cingulate Cortex ◽  
Entire Body ◽  
Mechanical Stimuli ◽  
Noxious Heat ◽  
Nociceptive Neurons

1. Single-unit responses in area 24 of cingulate cortex were examined in halothane-anesthetized rabbits during stimulation of the skin with transcutaneous electrical (TCES, 3-10 mA), mechanical (smooth or serrated forceps to the dorsal body surface or graded pressures of 100-1,500 g to the stabilized ear) and thermal (> 25 degrees C) stimulation. 2. Of 542 units tested in cingulate cortex, 150 responded to noxious TCES (> or = 6 mA), 93 of 221 units tested responded to noxious mechanical (serrated forceps) and 9 of 47 units tested responded to noxious heat (> 43 degrees C) stimuli. Twenty-five percent of the units that responded to noxious mechanical stimuli also responded to noxious heat stimuli. The only innocuous stimulus that evoked activity in cingulate cortex was a "tap" to the skin and this was effective for 11 of 14 tested units. 3. In 74 units that produced excitatory responses to TCES of the contralateral ear, response latency was 166 +/- 11.3 (SE) ms and response duration was 519 +/- 52.1 ms. 4. Twenty of the 150 units that responded to noxious TCES were initially inhibited. These responses were usually < 1 s in duration (17 of 20 units), whereas responses in the other 3 lasted for over 20 s. 5. Most units had broad receptive fields, because noxious mechanical stimuli anywhere on the dorsal surface of the rabbits, including the face and ears, evoked responses. A small number of units for which the entire body surface was tested (3 of 15 units) had receptive fields limited to the ears, rostral back, and forepaws. 6. Fifteen of 33 units tested had no preferential responses to noxious TCES of the ipsilateral and contralateral ears. Of the remaining units, 10 had a greater response to contralateral and 8 had a greater response to ipsilateral stimuli. 7. The locations of 186 units were histologically verified. Most nociceptive cingulate units were in dorsal area 24b in layers III (n = 35), II (n = 13), or V (n = 9). 8. Cortical knifecut lesions were made in five rabbits to determine if the responses in area 24 were dependent on lateral or posterior cortical inputs. These lesions did not alter the percentage of units driven by noxious stimuli nor response latency. 9. Injections of lidocaine were made into medial parts of the thalamus in six animals and injection and recording sites analyzed histologically.(ABSTRACT TRUNCATED AT 400 WORDS)

Responses of spinal trigeminal neurons to noxious stimulation of paranasal cavities – a rat model of rhinosinusitis headache

Cephalalgia ◽  
2020 ◽  
pp. 033310242097046
Author(s):  
Michael Koch ◽  
Julika Sertel-Nakajima ◽  
Karl Messlinger
Keyword(s):  
Potassium Chloride ◽  
Dura Mater ◽  
Paranasal Sinuses ◽  
Interstitial Fluid ◽  
Receptive Fields ◽  
Nasal Bone ◽  
Afferent Input ◽  
Noxious Stimulation ◽  
Trigeminal Neurons ◽  
Stimulation Of

Background The pathophysiology of headaches associated with rhinosinusitis is poorly known. Since the generation of headaches is thought to be linked to the activation of intracranial afferents, we used an animal model to characterise spinal trigeminal neurons with nociceptive input from the dura mater and paranasal sinuses. Methods In isoflurane anaesthetised rats, extracellular recordings were made from neurons in the spinal trigeminal nucleus with afferent input from the exposed frontal dura mater. Dural and facial receptive fields were mapped and the paranasal cavities below the thinned nasal bone were stimulated by sequential application of synthetic interstitial fluid, 40 mM potassium chloride, 100 µM bradykinin, 1% ethanol (vehicle) and 100 µm capsaicin. Results Twenty-five neurons with input from the frontal dura mater and responses to chemical stimulation of the paranasal cavities were identified. Some of these neurons had additional receptive fields in the parietal dura, most of them in the face. The administration of synthetic interstitial fluid, potassium chloride and ethanol was not followed by significant changes in activity, but bradykinin provoked a cluster of action potentials in 20 and capsaicin in 23 neurons. Conclusion Specific spinal trigeminal neurons with afferent input from the cranial dura mater respond to stimulation of paranasal cavities with noxious agents like bradykinin and capsaicin. This pattern of activation may be due to convergent input of trigeminal afferents that innervate dura mater and nasal cavities and project to spinal trigeminal neurons, which could explain the genesis of headaches due to disorders of paranasal sinuses.

Response and receptive-field properties of spinomesencephalic tract cells in the cat

1986 ◽  
Vol 55 (1) ◽  
pp. 76-96 ◽  
Author(s):  
R. P. Yezierski ◽  
R. H. Schwartz
Keyword(s):  
Spinal Cord ◽  
Receptive Field ◽  
Dynamic Range ◽  
Receptive Fields ◽  
The Body ◽  
Noxious Stimulation ◽  
Primary Afferent ◽  
Receptive Field Properties ◽  
Afferent Fibers ◽  
Primary Afferent Fibers

Recordings were made from 90 identified spinomesencephalic tract (SMT) cells in the lumbosacral spinal cord of cats anesthetized with alpha-chloralose and pentobarbital sodium. Recording sites were located in laminae I-VIII. Antidromic stimulation sites were located in different regions of the rostral and caudal midbrain including the periaqueductal gray, midbrain reticular formation, and the deep layers of the superior colliculus. Twelve SMT cells were antidromically activated from more than one midbrain level or from sites in the medial thalamus. The mean conduction velocity for the population of cells sampled was 45.2 +/- 21.4 m/s. Cells were categorized based on their responses to graded intensities of mechanical stimuli and the location of excitatory and/or inhibitory receptive fields. Four major categories of cells were encountered: wide dynamic range (WDR); high threshold (HT); deep/tap; and nonresponsive. WDR and HT cells had excitatory and/or inhibitory receptive fields restricted to the ipsilateral hindlimb or extending to other parts of the body including the tail, forelimbs, and face. Some cells had long afterdischarges following noxious stimulation, whereas others had high rates of background activity that was depressed by nonnoxious and noxious stimuli. Deep/tap cells received convergent input from muscle, joint, or visceral primary afferent fibers. The placement of mechanical lesions at different rostrocaudal levels of the cervical spinal cord provided information related to the spinal trajectory of SMT axons. Six axons were located contralateral to the recording electrode in the ventrolateral/medial or lateral funiculi while two were located in the ventrolateral funiculus of the ipsilateral spinal cord. Stimulation at sites used to antidromically activate SMT cells resulted in the inhibition of background and evoked responses for 22 of 25 cells tested. Inhibitory effects were observed on responses evoked by low/high intensity cutaneous stimuli and by the activation of joint or muscle primary afferent fibers. Based on the response and receptive-field properties of SMT cells it is suggested that the SMT may have an important role in somatosensory mechanisms, particularly those related to nociception.

Extrasynaptic receptors on cell bodies of neurons in central nervous system of the leech

1977 ◽  
Vol 40 (2) ◽  
pp. 446-452 ◽  
Author(s):  
P. B. Sargent ◽  
K. W. Yau ◽  
J. G. Nicholls
Keyword(s):  
Central Nervous System ◽  
Nervous System ◽  
Glutamic Acid ◽  
Systematic Study ◽  
Receptive Fields ◽  
Aminobutyric Acid ◽  
Gamma Aminobutyric Acid ◽  
Light Touch ◽  
Nociceptive Neurons ◽  
Extrasynaptic Receptors

1. A systematic study has been made of the sensitivity of identified sensory and motoneurons in the leech central nervous system to chemical transmitter substances. 2. The following substances elicited responses from the cell bodies of individual neurons: acetylcholine, 5-hydroxytryptamine, gamma-aminobutyric acid, glutamic acid, glycine, dopamine, and norepinephrine. Since the cell bodies of leech neurons are free of synapses, the receptors that give rise to these responses are extrasynaptic. 3. Sensory and motoneurons of different function had characteristic complements of extrasynaptic receptors. For example, mechanosensory cells responding to light touch, to pressure, and to noxious stimuli could be distinguished by their responses to iontophoretically applied compounds. For one of these modalities (nociceptive), neurons with different receptive fields but otherwise similar properties had markedly distinct extrasynaptic receptors. The possible significance of extrasynaptic receptors is discussed.

Leech Mechanosensation

2017 ◽  
Author(s):  
Brian D. Burrell
Keyword(s):  
Central Nervous System ◽  
Nervous System ◽  
Sensory Neurons ◽  
Receptive Fields ◽  
Model Systems ◽  
Sensory Cells ◽  
Mechanical Stimuli ◽  
Nociceptive Neurons ◽  
Mechanosensory Neurons ◽  
The Central Nervous System

The medicinal leech (Hirudo verbana) is an annelid (segmented worm) and one of the classic model systems in neuroscience. It has been used in research for over 50 years and was one of the first animals in which intracellular recordings of mechanosensory neurons were carried out. Remarkably, the leech has three main classes of mechanosensory neurons that exhibit many of the same properties found in vertebrates. The most sensitive of these neurons are the touch cells, which are rapidly adapting neurons that detect low-intensity mechanical stimuli. Next are the pressure cells, which are slow-adapting sensory neurons that respond to higher intensity, sustained mechanostimulation. Finally, there are nociceptive neurons, which have the highest threshold and respond to potentially damaging mechanostimuli, such as a pinch. As observed in mammals, the leech has separate mechanosensitive and polymodal nociceptors, the latter responding to mechanical, thermal, and chemical stimuli. The cell bodies for all three types of mechanosensitive neurons are found in the central nervous system where they are arranged as bilateral pairs. Each neuron extends processes to the skin where they form discrete receptive fields. In the touch and pressure cells, these receptive fields are arranged along the dorsal-ventral axis. For the mechano-only and polymodal nociceptive neurons, the peripheral receptive fields overlap with the mechano-only nociceptor, which also innervates the gut. The leech also has a type of mechanosensitive cell located in the periphery that responds to vibrations in the water and is used, in part, to detect potential prey nearby. In the central nervous system, the touch, pressure, and nociceptive cells all form synaptic connections with a variety of motor neurons, interneurons, and even each other, using glutamate as the neurotransmitter. Synaptic transmission by these cells can be modulated by a variety of activity-dependent processes as well as the influence of neuromodulatory transmitters, such as serotonin. The output of these sensory neurons can also be modulated by conduction block, a process in which action potentials fail to propagate to all the synaptic release sites, decreasing synaptic output. Activity in these sensory neurons leads to the initiation of a number of different motor behaviors involved in locomotion, such as swimming and crawling, as well as behaviors designed to recoil from aversive/noxious stimuli, such as local bending and shortening. In the case of local bending, the leech is able to bend in the appropriate direction away from the offending stimuli. It does so through a combination of which mechanosensory cell receptive fields have been activated and the relative activation of multiple sensory cells decoded by a layer of downstream interneurons.

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