Journal of Neurophysiology

Janig, W.; Koltzenburg, M.

doi: N/Apmid: 1869905

Abstract 1. Conscious perception of noxious and innocuous distension of the colon as well as the reflex control of anal continence and defecation largely depend on an intact sacral primary afferent innervation. Here we have studied the functional properties of these visceral primary afferent neurons in the dorsal root S2 in 17 cats. Single fibers projecting into the pelvic nerve were identified electrically and studied with innocuous and noxious mechanical stimulation of colon and anal canal. 2. A total of 59 units responding to one of these stimuli were investigated and they could be separated into two subpopulations of afferents. Thirty-six fibers were reproducibly excited by distension of the colon, but not by mechanical stimulation of the anal canal. They were thin myelinated or unmyelinated fibers with a median conduction velocity of 3.2 m/s. The remaining 23 units had receptive fields in the mucosa of the anal canal and responded readily to an innocuous proximodistal shearing stimulus, but not to distension stimuli applied to the same area. All, but two of these afferents were thin myelinated with a median conduction velocity of 7.7 m/s, which was significantly different from the conduction velocity of afferent neurons responding to distension of the colon. 3. Units responding to distension of the colon had thresholds in the innocuous range of the intracolonic pressure. Receptors that were activated only by noxious intraluminal pressure were absent. On the basis of their response to supramaximal isotonic distension, colonic afferents could be subclassified as phasic (n = 17) or tonic (n = 19) units. Phasic afferents were only transiently excited during filling or emptying of the colon, whereas tonic afferents discharged throughout the distension. The two populations had also significantly different median conduction velocities of 8.0 (n = 16) and 1.7 (n = 15) m/s, respectively. 4. Stimulation response functions were evaluated for 12 tonic afferents. All units encoded an increase of intracolonic pressure by the intensity of their discharge frequency. Increases of intracolonic pressure produced significantly higher discharge frequencies from unmyelinated than from thin myelinated afferents. 5. In three animals the percentage of unmyelinated fibers responding to mechanical stimulation of colon and anal canal was determined. Out of 213 electrically identified unmyelinated units projecting into the pelvic nerve, only 11 (5.2%) were excited. Thus, acute innocuous and noxious mechanical stimuli of the large intestine do not appear to be the adequate stimulus for the large majority of unmyelinated pelvic afferents. 6. In conclusion, distension of the colon and mechanical stimulation of the anal canal activates distinct populations of primary afferent neurons.(ABSTRACT TRUNCATED AT 400 WORDS) Copyright © 1991 the American Physiological Society

Smith, D. V.; Hanamori, T.

doi: N/Apmid: 1869907

Abstract 1. Mammalian taste receptors are distributed within several distinct subpopulations, innervated by branches of cranial nerves VII, IX, and X. Most gustatory electrophysiology has focused on input from the fungiform papillae on the anterior portion of the tongue, carried by the chorda tympani branch of the VIIth nerve. However, laryngeal taste buds in the hamster are as numerous as those in the fungiform papillae. Gustatory fibers in the hamster's chorda tympani and glossopharyngeal nerves have been well characterized. In comparison with these taste fibers, much less is known about the chemical sensitivities of fibers innervating laryngeal taste buds. 2. Action potentials were recorded from 65 individual fibers in the superior laryngeal nerve (SLN) of the hamster. Stimuli were distilled H2O and five concentrations each of sucrose, NaCl, HCl, and quinine hydrochloride (QHCl). All stimuli except the NaCl series were made in physiological saline (0.154 M NaCl) and were delivered from the laryngeal side of the epiglottis via a tracheal cannula. Responses were quantified as the number of impulses in 10 s minus the responses in the preceding 10 s of baseline activity during a rinse with physiological saline. 3. Distilled H2O, HCl, and NaCl were by far the most excitatory stimuli, with mean responses across all cells 5-10 times greater than those evoked by sucrose or QHCl. The order of effectiveness of the strongest concentrations of the stimuli was H2O greater than 0.03 M HCl greater than 1.0 M NaCl much greater than 0.03 M QHCl greater than 1.0 M sucrose. 4. The mean concentration-response function for NaCl was U shaped, with the greatest number of impulses to distilled H2O and 1.0 M NaCl. The responses diminished as the concentrations approached physiological levels (0.154 M NaCl), where there was no response, and increased as NaCl concentration rose above this level. Increasing concentrations of HCl above 0.0003 M elicited increasing responses in these fibers. 5. The mean time course of the responses to distilled H2O and to hypotonic NaCl solutions (0.01 and 0.03 M) peaked in the first few seconds and then declined slowly. This was distinct from the time course of the responses to hypertonic NaCl concentrations (0.3 and 1.0 M), which increased gradually throughout the 10-s response period. Responses to HCl peaked in the initial second and then decayed rapidly to a slowly declining plateau. These distinctively different time courses suggest different receptor mechanisms for water, salt, and acid stimuli. 6. The across-fiber pattern of the responses to hypotonic NaCl solutions correlated strongly to that elicited by distilled H2O.(ABSTRACT TRUNCATED AT 400 WORDS) Copyright © 1991 the American Physiological Society

Aram, J. A.; Michelson, H. B.; Wong, R. K.

doi: N/Apmid: 1678421

Abstract 1. Intracellular and extracellular recordings were carried out in guinea pig neocortical slices to examine the effects of blockade of excitatory amino acid (EAA) synaptic transmission on population discharges elicited by 4-aminopyridine (4-AP; 50-100 microM). 2. After the introduction of 4-AP, two distinct types of rhythmic spontaneous field potentials were recorded in neocortical slices. Type I consisted of multiple spike discharges lasting 20-90 s. These events occurred at a frequency of 0.4-0.2/min. Type II were single field potential spikes (3-6 s in duration) occurring at a higher frequency (2-4/min). 3. Blockade of amino acid-mediated excitatory synaptic transmission with D-2-amino-5-phosphonovaleric acid (D-AP5; 10-30 microM) or 3-(2-carboxypiperazin-4-yl)propyl-l-phosphonic acid (CPP, 10 microM) and 6-cyano-7nitroquinoxaline-2,3-dione (CNQX; 10 microM) abolished the first type of 4-AP-induced field potential, whereas type II events persisted. 4. Type II field events, occurring in the presence of EAA blockers, were further characterized by paired recordings. Events recorded along an axis orthogonal to the pia surface occurred simultaneously without measurable delay. Recordings made along a plane parallel to the pia surface showed that type II discharges propagated over distances of greater than or equal to 3 mm at an estimated velocity of 7.5 mm/s. 5. Intracellular recordings show that during type II field discharges all cells exhibited phasic depolarizations or hyperpolarizations, depending on the resting membrane potential. When resting potentials were more depolarized than -68 mV, events became mostly hyperpolarizing.(ABSTRACT TRUNCATED AT 250 WORDS) Copyright © 1991 the American Physiological Society

Robertson, R. M.

doi: N/Apmid: 1869910

Abstract 1. Synaptic interactions between identified neurons in the flight system of the locust were investigated by the use of standard intracellular recording and staining techniques. The intent was to determine the distribution and functional significance of delayed excitatory connections, which have been previously described. 2. For one inhibitory connection it was demonstrated that subthreshold depolarization of the presynaptic neuron was sufficient to cause release of transmitter at the synapse. This established the existence of graded interactions between spiking flight neurons. 3. Three inhibitory interneurons were found to cause delayed excitatory responses in several other neurons. Often these were coupled with direct inhibitory connections between the same pre- and postsynaptic neurons, resulting in an inhibitory/excitatory (I/E) postsynaptic potential (PSP). The two phases of this PSP were variable. 4. Delayed excitatory connections appeared powerful while the flight system was inactive. However, these connections were disabled during flight rhythms at the phase when the presynaptic neuron was depolarized and firing action potentials. This was likely to be due to the nature of the disynaptic disinhibitory interaction being via (an) intervening neuron(s) with oscillating membrane potentials and thresholds for release of transmitter. 5. Thus connections demonstrated when flight rhythms were not expressed changed their character during flight rhythms. The delayed excitatory connections in this system probably reflect complex circuits of inhibition mediated by graded interactions and have little functional significance as phenomena in their own right. Copyright © 1991 the American Physiological Society

Wu, S. M.

doi: N/Apmid: 1651374

Abstract 1. The input-output relation of the feedback synapse between horizontal cells (HCs) and cones was studied by simultaneously recording the light responses of the HCs and of cones the outer segments of which were truncated off. 2. The postsynaptic light response of the truncated cone was depolarizing and free of direct influence of photocurrents. These postsynaptic light responses were graded and sustained; their waveform resembled that of the HC light responses. 3. Input-output relation of the HC-cone feedback synapse was obtained by plotting the simultaneous voltage points of the HC and truncated cone light responses. At the resting potential of the cone (-40 mV), the voltage gain of the feedback synapse was about -0.33 when VHC = -20 mV and it was about -0.03 when VHC = -60 mV. 4. At more hyperpolarized cone voltages, the feedback signals in cones became smaller, and they reversed at about -67 mV. The voltage gain of the feedback synapse at VHC = -20 mV was about -0.23, -0.18, -0.07, and +0.2 when Vcone = -44.5, -52.5, -65, and -77.5 mV, respectively. 5. Light hyperpolarized the HC, which resulted in a conductance change (delta Gs) in cones. The cone conductance decreased progressively as the HCs were increasingly hyperpolarized, and delta Gs reached a maximum value of 0.93 nS when the HCs were hyperpolarized from -20 to -52 mV. 6. The peak light responses of intact cones were plotted against the peak HC light responses. This gives the relationship between the pre- and postsynaptic voltages of the cone-HC (forward) and HC-cone (feedback) synapses at any given light intensity. Combining this relationship with the input-output relations obtained at various voltages of the truncated cones allows the input-output relation of the feedback synapse for light-evoked signals to be obtained. 7. The input-output relation of the feedback synapse for light-evoked signals was bell-shaped, because the feedback light responses were controlled by two opposing factors: as the light became brighter, the postsynaptic conductance change increased, but the driving force decreased. 8. For light-evoked signals, the slope gain (slope of the input-output relation) of the HC-cone feedback synapse was negative (varied from -0.33 to 0) when VHC lay between -20 and -52 mV; and it was positive (0 to +0.8) when VHC lay between -52 and -72 mV. 9.(ABSTRACT TRUNCATED AT 400 WORDS) Copyright © 1991 the American Physiological Society

Paige, G. D.; Tomko, D. L.

doi: N/Apmid: 1869911

Abstract 1. The purpose of this study was to quantify the response characteristics of eye movements produced by linear head oscillations in the dark (the linear vestibuloocular reflex, or LVOR). Horizontal, vertical, and torsional eye movements were measured in adult squirrel monkeys by the use of a dual scleral search-coil technique during linear oscillations (0.5, 1.5, and 5.0 Hz, 0.36 g peak acceleration) along the animals' interaural (IA), dorsoventral (DV), and nasooccipital (NO) axes. 2. Two LVOR responses, horizontal eye movements during IA-axis translation and vertical eye movements during DV-axis motion, were in a compensatory direction for head translation. Response amplitudes increased as frequency increased, whereas phase typically showed a lead. 3. Two other LVORs, torsional responses during IA-axis translation (all frequencies) and vertical responses during NO-axis oscillations (0.5 Hz), behaved differently. These two LVORs cannot be functionally compensatory for head translation because they degrade fixation on targets, and therefore image stability, by rotating the eyes off target (NO-vertical) or torting the eyes relative to the visual world (IA-torsional). Responses to NO-axis motion at frequencies greater than 0.5 Hz depended on initial eye position and fixation distance and are described in the companion paper. 4. The effect of head orientation on the LVOR was assessed by testing four head positions in 90 degrees steps around the axis of head motion for each of the three axes of translation. This was done, first, to determine whether the LVORs are responses to the "swinging vector" of gravitoinertial force during linear head motion or to head translation; and second, to quantify potential effects of static head (otolith) orientation on the LVORs. Results showed no systematic effects of head orientation on LVOR responses in the frequency bandwidth studied. This indicates that the LVORs are dependent on the direction of linear motion relative to the head (and otolith organs) but not on the swinging vector of gravitoinertial force, and that the LVORs are uninfluenced by static orientation of the head and reloading of the otoliths. Copyright © 1991 the American Physiological Society

Paige, G. D.; Tomko, D. L.

doi: N/Apmid: 1869912

Abstract 1. Horizontal, vertical, and torsional eye movements were recorded (search coil technique) from five squirrel monkeys during horizontal linear oscillations at 0.5, 1.5, and 5.0 Hz, 0.36 g peak acceleration. Monkeys were positioned to produce linear motion in their nasooccipital (NO), interaural (IA), and dorsoventral (DV) axes. Responses of the linear vestibuloocular reflex (LVOR) were recorded in darkness and in the light with the subjects viewing a head-fixed field 22 or 9.2 cm from the eye. The latter condition provided a measure of "visual suppression" of the LVOR (VSLVOR). Responses were also recorded while monkeys viewed earth-fixed targets, which allowed visual enhancement of the LVOR (VLVOR). Vergence angle was recorded in two monkeys to assess directly the point of binocular fixation in space during linear motion. 2. Two LVOR response types, vertical responses during 0.5-Hz NO-axis translation (NO-vertical) and torsional responses at all frequencies during IA-axis oscillation (IA-torsional) could not be compensatory reflexes for head translation because they either move the eye off target (NO-vertical) or tort the eye relative to the visual world (IA-torsional), thereby degrading visual image stability. 3. Other response types are considered compensatory because they help maintain ocular fixation in space during linear head translation. These include horizontal responses to IA-axis motion (IA-horizontal), vertical responses to DV-axis translation (DV-vertical), and both horizontal and vertical responses to NO-axis oscillation (1.5 and 5 Hz). Observations focus on responses to 5-Hz oscillations, in which visual inputs are essentially ineffective in modifying the LVOR. 4. The kinematics of perfect ocular compensation during head translation indicate that the ideal ocular response is governed by the motion of the eye relative to target position. Relevant variables include target distance, which is crucial for all axes of motion, and target eccentricity, which is important only for head motion roughly parallel to the target (NO-axis translation). Findings are compatible with predictions based on ideal kinematics. However, it is the point of binocular fixation in space, not actual target position, that governs LVOR behavior. 5. The IA-horizontal and DV-vertical LVOR is in response to head motion roughly orthogonal to the line of sight. Responses under all stimulus conditions (LVOR, VSLVOR, and VLVOR) behaved similarly at 5 Hz, and were modulated linearly with vergence in meter angles (MA), the reciprocal of binocular fixation distance.(ABSTRACT TRUNCATED AT 400 WORDS) Copyright © 1991 the American Physiological Society

Noth, J.; Schwarz, M.; Podoll, K.; Motamedi, F.

doi: N/Apmid: 1831227

Abstract 1. The aim of the present study was to identify the type of spinal afferents involved in the generation of the long-latency response in intrinsic human hand muscles. Position-controlled extensions were imposed on the index finger or on the wrist of healthy subjects who were exerting a steady voluntary flexion force at the relevant joint. Averaged surface electromyographic (EMG) responses of the first dorsal interosseus muscle (FDI) or of the wrist flexors were evaluated with respect to latency and size. 2. Small transient angular displacements of the index finger (1 degree, as measured at the metacarpophalangeal joint), which are supposed to excite primary rather than secondary afferents, evoked two clearly discernible EMG responses with mean latencies of 32.3 ms (M1 response) and 54.7 ms (M2 response), respectively. The size of the M2 response exceeded the size of the M1 response by 60%. In the wrist flexors, transient stretch (1 degree) gave rise to a large M1 response (latency 22.8 ms) and a small, inconstent M2 response. 3. Small-amplitude vibration of the index finger elicited EMG responses in the FDI that were qualitatively and quantitatively similar to those seen in response to small transient stretches of the index finger. This was also true for fast ramp-and-hold stretches (stretch velocity 400 degrees/s, amplitude 5 degrees), whereas slow ramp-and-hold stretches (125 degrees/s, 5 degrees) elicited predominantly M2 responses. 4. In the FDI, the mechanical threshold of the M1 and M2 response to the transient angular displacement was approximately 0.15 degrees, with a tendency for the M2 response to appear at a lower threshold.(ABSTRACT TRUNCATED AT 250 WORDS) Copyright © 1991 the American Physiological Society

Fyffe, R. E.

doi: N/Apmid: 1869909

Abstract 1. Intracellular staining of Renshaw cells and alpha motoneurons was used to determine the spatial distribution of recurrent inhibitory synapses on spinal motoneurons in the cat. In each experiment, a Renshaw cell and one or more possible target motoneurons were labeled with horseradish peroxidase after physiological identification. 2. Paris of labeled neurons were reconstructed and measured at the light microscopic level. As defined by light microscopy, presumed synaptic contacts between nine Renshaw cells and 10 postsynaptic motoneurons were observed. On average, each Renshaw cell made three synaptic contacts (range 1-9) on each motoneuron. 3. Electron microscopic confirmation of several presumed contacts provided evidence that the appositions identified by light microscopic criteria are genuine contacts between Renshaw cell boutons and the labeled motoneuron. 4. All of the identified synapses observed in these experiments were located on motoneuron dendrites, between 65 and 706 microns from the soma. Use of a simplified cable model indicated that the synapses are electrotonically close to the soma, the average location being approximately 0.25 length constants from the soma (range 0.04-0.82 lambda). 5. These observations provide direct evidence to support the hypothesis that Renshaw cell synapses on motoneurons are located on the dendrites and not on the cell body (whereas reciprocal inhibitory synapses, from Ia inhibitory interneurons, are predominantly located on the soma). The functional significance of the observed distribution of Renshaw inhibitory synapses is discussed. One possibility is that the recurrent inhibitory pathway selectively inhibits particular dendritic inputs. Copyright © 1991 the American Physiological Society

Rosenberg, A. F.; Ariel, M.

doi: N/Apmid: 1869903

Abstract 1. The direct retinal input pathway to the basal optic nucleus (BON), the primary nucleus of the turtle accessory optic system, was characterized physiologically. We tested the hypothesis that directional information encoded in retinal ganglion cells can influence the BON via a direct pathway. Using an in vitro whole-brain, eyes-attached preparation, we demonstrated the directness of this pathway by 1) antidromic activation of retinal ganglion cells from the contralateral BON and 2) orthodromic activation of the BON from the contralateral optic nerve. 2. Of 72 physiologically classified retinal ganglion cells, 9 could be antidromically activated from the contralateral BON with low current (less than 200 micro A). Eight of these cells were direction-sensitive (DS). The ninth cell did not respond to visual stimulus movement. The antidromic latencies ranged from 2.2 to 6.1 ms with a mean of 3.8 ms. These latencies were quite consistent for each cell, having an average SD of 0.08 ms. Moreover, consistent responses could always be recorded at stimulation rates up to 100 Hz. 3. With current stimulation of the contralateral optic nerve, the orthodromic conduction latency of 17 BON single units ranged from 2.5 to 6.6 ms with a mean of 4.6 ms. These latencies were more variable for an individual cell, having an average SD of 0.3 ms. Responses to individual current pulses could never be consistently evoked at stimulation rates greater than 40 Hz. 4. DS responses were recorded in BON single units after the removal of the dorsal midbrain, including the optic tectum and pretectum as well as the telencephalon. Three of these cells were activated orthodromically by current stimulation delivered to the contralateral optic nerve. Thus directional information reaches the BON via a direct projection from the contralateral retina. 5. Visual response properties of DS retinal ganglion cells were compared with those of BON cells to examine the transformations that take place in the brain stem. Applying a limacon model to the responses of both DS retinal ganglion cells and BON cells revealed that both types of cells have very similar direction tuning. However, the distribution of maximally responsive directions in the retina may differ from that of the BON. 6. Because DS retinal ganglion cells project directly to the BON, and because BON cells lose their direction sensitivity after retinal application of GABA antagonists, we conclude that the BON receives essential directional information directly from DS retinal ganglion cells. This directional information in the BON may represent a retinal slip error signal necessary for retinal image stabilization. Copyright © 1991 the American Physiological Society