Contribution of antidromic efferent-fiber excitation to mass spinal potentials in the cat

The contribution of antidromic excitation of motoneurons to cord dorsum potentials (CDP) was studied in the spinal cord of anesthetized cats. It was shown that stimulation of ventral roots (VR) or peripheral nerves following deafferentiation of a number of segments by crosscutting of dorsal roots on the dorsal surface evokes appreciable positive-negative CDP (VR-CDP). Under intact conditions, VR effects of antidromic stimulation of efferent fibers brings appreciable input to the initial "fast" CDP component (the "afferent" peak); input values for the main mixed nerves of the hindlimb are presented. After conditioning stimulation of a mixed nerve, VR-CDP undergo inhibition with two maximums, associated with blocking of the effects of antidromic excitation of efferents by orthodromic mono- and polysynaptic reflex discharges of motoneurons. The hypothesis that intactness of efferents in nerves under stimulation can be determined from an analysis of initial CDP components is stated.Scientific-Research Institute of Biology, Dnepropetrovsk State University, Dnepropetrovsk. Translated from Neirofiziologiya, Vol. 23, No. 6, pp. 655–661, November–December, 1991.     

Antidromic vasodilation caused by A-delta afferents in the frog

In anesthetized immobilized frog we recorded changes in hind leg volume evoked by electrical stimulation of peripheral end of the sciatic nerve. The ranges of the stimulus amplitudes sufficient to induce vasodilator or vasoconstrictor reactions were estimated. In a separate set of experiments thresholds of A alpha beta, A delta and C-afferent fibers excitation were evaluated by recording waves of compound action potentials in VIII-X dorsal roots. It was found that vasodilation is elicited by the stimuli of virtually the same intensity range as the excitation of A delta afferent fibers, including low threshold one. Consequently we concluded that in frog these myelinated afferent fibers are capable of dilating the blood vessels by antidromic action. This finding is in contrast with antidromic vasodilation in mammals which is known to result mainly from the impulses of the unmyelinated afferent fibers.     

Impulse Origin and Propagation in a Bipolar Sensory Neuron

Intracellular recording techniques were used to study electrical activity in bipolar sensory cells associated with crayfish tactile receptors. Several lines of evidence indicate that spikes evoked by natural stimulation of the receptor originate at a dendritic locus. Although overshooting spikes are recorded in the soma in response to both natural and antidromic stimulation receptor potentials are observed only rarely, and, when present, their amplitude is less than 5 mv. Impulses propagating centrifugally into the soma following antidromic stimulation always exhibit an inflection in the rising phase of the spike; however, orthodromic spikes are usually uninflected. Occasionally, orthodromic responses (in the soma) exhibit rather unusual wave forms. Such spikes evoked by natural stimuli are indistinguishable from those elicited electrically in the dendrite, but they do not resemble antidromic impulses. Because the axonal and dendritic boundaries of the soma have a low safety factor for spike transmission, at high frequencies invasion of the soma by dendritic spikes is impeded and often blocked. The soma region can thus act as a low-pass filter. The significance of this self-limiting mechanism for the behavior of the animal is not known; it is suggested, however, that this impediment is a potentially critical one, and may, in other situations, have encouraged the evolution of alternative arrangements.     

Intersegmental interneurons serving larval and pupal mechanosensory reflexes in the moth Manduca sexta

1. Intersegmental interneurons (INs) that participate in the larval bending reflex and the pupal gin trap closure reflex were identified in the isolated ventral nerve cord of Manduca sexta. INs 305, 504, and 703 show qualitatively different responses in the pupa than in the larva to electrical stimulation of sensory neurons that are retained during the larval-pupal transition to serve both reflexes. Action potentials produced by current injected into the 3 interneurons excite motor neurons that are directly involved in the larval and pupal reflexes. The excitation of the motor neurons is not associated with EPSPs at a fixed latency following action potentials in the interneurons, and thus there do not seem to be direct synaptic connections between the interneurons and the motor neurons. 2. IN 305 (Fig. 2) has a lateral soma, processes in most of the dorsal neuropil ipsilateral to the soma, and a crossing neurite that gives rise to a single contralateral descending axon. IN 305 is excited by stimulation of the sensory nerve ipsilateral to its soma in the larva and the pupa. Stimulation of the sensory nerve contralateral to its soma produces an inhibitory response in the larva, but a mixed excitatory/inhibitory response to the identical stimulus in the pupa. 3. IN 504 (Fig. 3) has a lateral soma, processes throughout most of the neuropil ipsilateral to the soma, and a crossing neurite that bifurcates to give rise to a process extending to the caudal limit of the neuropil and an ascending axon. IN 504 is excited by stimulation of the sensory nerve ipsilateral to its soma in both larvae and pupae, while the response to stimulation of the sensory nerve contralateral to its soma is inhibitory in the larva but mixed (excitatory/inhibitory) in the pupa. 4. IN 703 has a large antero-lateral soma, a neurite that extends across to the contralateral side giving rise to processes located primarily dorsally in both ipsilateral and contralateral neuropils, and two axons that ascend and descend in the connectives contralateral to the soma (Fig. 4). IN 703 responds to stimulation of the sensory nerves on either side of the ganglion, but the form of the response changes during the larval-pupal transition. In the larva, the response consists of very phasic (0-2 spikes) excitation, but in the pupa there is a prolonged excitation that greatly outlasts the stimulus (Fig. 6).(ABSTRACT TRUNCATED AT 400 WORDS)     

The innervation of the wind-sensitive head hairs of the locust, Schistocerca gregaria

ABSTRACT. The innervation of the locust head hairs was investigated by filling the sub-cuticular neurones with cobalt and by recording antidromic spikes at individual hairs when the circumoesophageal and cervical connectives were stimulated. The central projections from most head hairs ( c. 80%) terminate in the suboesophageal ganglion, whereas about 20% extend into the thoracic ganglia. Sensilla with projections to the thoracic ganglia are structurally no different from those whose fibres terminate in the suboesophageal ganglion and they are scattered throughout the hair fields with no consistent positions in different individuals. We have re-investigated the 'accessory response' (Camhi, 1969) and conclude that it is not indicative of a separate afferent or efferent system but is simply cross-talk from adjacent axons. We relate our findings to the fibre composition of the dorsal tegumentary nerve.     

Steps in the production of motoneuron spikes

1. Spikes evoked in spinal motoneurons by antidromic stimulation normally present an inflection in their rising phase. A similar inflection is present in spikes evoked by direct stimulation with short pulses. 2. In either case the inflection becomes less prominent if the motoneuron membrane is depolarized and more prominent when it is hyperpolarized. Both antidromic and direct spikes may fall from the level of the inflection thus evoking a "small spike" only if sufficient hyperpolarization is applied. Similar events occur when antidromic or direct spikes are evoked in the aftermath of a preceding spike. 3. Spikes evoked by direct stimuli applied shortly after firing of a "small spike" may also become partially blocked at a critical stimulus interval. At shorter intervals, however, spike size again increases and no inflection can be detected in the rising phase. 4. When a weak direct stimulus evokes a small spike only, a stronger stimulus may evoke a full spike. Curves of the strength of the stimuli required for eliciting small or full spikes have been constructed in a number of conditions. 5. To explain the results it is assumed that threshold of the major portions of the soma membrane is higher than the threshold of the axon, the transition occurring over a finite area near the axon hillock. Following antidromic or direct stimulation, soma excitation is then initiated in the region of the axon hillock. Spread of activity towards the soma occurs at first slowly and with low safety factor. At this stage block may be easily evoked. Safety factor for propagation increases rapidly as the growing impulse involves larger and larger areas of the soma membrane so that, once the critical areas are excited, activation of the remaining portions of the soma membrane will suddenly occur.     

Antidromic discharges of dorsal root afferents in the neonatal rat.

Presynaptic inhibition of primary afferents can be evoked from at least three sources in the adult animal: 1) by stimulation of several supraspinal structures; 2) by spinal reflex action from sensory inputs; or 3) by the activity of spinal locomotor networks. The depolarisation in the intraspinal afferent terminals which is due, at least partly, to the activation of GABA(A) receptors may be large enough to reach firing threshold and evoke action potentials that are antidromically conducted into peripheral nerves. Little is known about the development of presynaptic inhibition and its supraspinal control during ontogeny. This article, reviewing recent experiments performed on the in vitro brainstem/spinal cord preparation of the neonatal rat, demonstrates that a similar organisation is present, to some extent, in the new-born rat. A spontaneous activity consisting of antidromic discharges can be recorded from lumbar dorsal roots. The discharges are generated by the underlying afferent terminal depolarizations reaching firing threshold. The number of antidromic action potentials increases significantly in saline solution with chloride concentration reduced to 50% of control. Bath application of the GABA(A) receptor antagonist, bicuculline (5-10 microM) blocks the antidromic discharges almost completely. Dorsal root discharges are therefore triggered by chloride-dependent GABA(A) receptor-mediated mechanisms; 1) activation of descending pathways by stimulation delivered to the ventral funiculus (VF) of the spinal cord at the C1 level; 2) activation of sensory inputs by stimulation of a neighbouring dorsal root; or 3) pharmacological activation of the central pattern generators for locomotion evokes antidromic discharges in dorsal roots. VF stimulation also inhibited the response to dorsal root stimulation. The time course of this inhibition overlapped with that of the dorsal root discharge suggesting that part of the inhibition of the monosynaptic reflex may be exerted at a presynaptic level. The existence of GABA(A) receptor-independent mechanisms and the roles of the antidromic discharges in the neonatal rat are discussed.     

Synaptic morphology of the carotid body of the domestic fowl

Efferent and reciprocal synapses have been demonstrated in the carotid body of the domestic fowl (Gallus gallus domesticus). Synapses were also found with purely afferent morphology, but were probably components of reciprocal synapses. The general morphology of the endings suggested the presence of two types of axon, afferent axons making reciprocal and perhaps afferent synapses with Type I cells, and efferent axons making efferent synapses with Type I cells. A few axo-dendritic synapses were also found. The dense-cored vesicles associated with the afferent components of reciprocal synapses and with the possible true afferent synapses varied in diameter and core but could belong to one population of pre-synaptic vesicles. These observations are consistent wtih a new theory for the carotid body receptor mechanism. This proposes a spontaneously discharging afferent axon inhibited by an inhibitory transmitter substance released by the Type I cell via the "afferent" component of its reciprocal synapse, the "efferent" component inhibiting this release. Besides this chemoreceptor modulation of its afferent axon, the Type I cell may also have a general secretory function.     

Depolarization of primary afferents during fictitious scratching in thalamic cats

The dorsal cord and dorsal root potentials were recorded in immobilized thalamic cats during fictitious scratching evoked by mechanical stimulation of the ear. Depolarization of primary afferents was shown to be simulated by the central scratching generator. Antidromic spike discharges appeared at the peak of the primary afferent depolarization waves in certain afferent fibers. Similar discharges arise in the resting state in response to stimulation of limb mechanoreceptors. It is suggested that during real scratching primary afferent depolarization and antidromic spikes evoked by it may effectively modulate the level of the afferent flow to spinal neurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 2, pp. 173–176, March–April, 1978.     

Site of Origin and Propagation of Spike in the Giant Neuron of Aplysia

The form and time sequence of spikes generated by orthodromic, antidromic, and direct stimulation and during spontaneous activity have been studied with intracellular electrodes simultaneously introduced in the soma and in different parts of the axon of the giant nerve cell of Aplysia. Evidence was obtained that under normal conditions of excitability, the spike originates at some distance from the soma in an axonal region with a higher excitability surpassing that of the surrounding membranes. Between the trigger zone and the soma is situated a region of transitional excitability where the conduction of the spike towards the soma may be blocked at a functionally determined and variable locus. The cell body is electrically excitable, but has the highest threshold of all parts of the neuron. The inactivation or even the removal of the cell body does not suppress synaptic transmission.     

Neural mechanisms underlying behavior in the locust Schistocerca gregaria I. Physiology of identified motorneurons in the metathoracic ganglion

A preparation of the desert locust, Schistocera gregaria, has been developed, in which it was possible to work with identified neurons while still allowing some behavior. A total of 26 motorneurons to the hind leg were studied singly, and in various pairs, both by direct stimulation, and by recording during spontaneous activity and various reflex actions. Motorneurons were identified by passing current into their somata and correlating the evoked somata spikes with extracellularly or intracellularly recorded events in the muscles. Tension of the muscle was also recorded and motor axons were stimulated to evoke antidromic spikes in the somata. Both epsp's and ipsp's can be seen clearly in recordings from the somata; spikes appear as electrotonically conducted remnants only. Somata exhibited little or no electrogenesis. It is inferred that impulses are initiated in a zone tentatively identified with the region of emergence of the motor axon from the neuropil. Integration occurs in the neuropilar segment, with the soma serving as a parallel RC element. Data was obtained on the central mechanisms of coordination of synergistic and antagonistic motorneurons and on the modes of excitation of slow and fast neurons to the same muscles.     

Modes of Initiation and Propagation of Spikes in the Branching Axons of Molluscan Central Neurons

A study has been made of Aplysia nerve cells, mainly in the pleural ganglia, in which the main axon divides into at least two branches in the neighbourhood of the soma. Conduction between these branches was investigated by intracellular recordings from the soma following antidromic stimulation via the nerves containing the axonal branches. It has been shown that transmission between separate branches need not involve discharge of the soma but only of the axonal region between the soma and the origin of the branches. In some cells, the spike may fail to invade the other axonal branch, whereas transmission in the opposite direction is readily achieved. Often spikes in none of the branches are transmitted to the others, unless facilitated. Indications about the geometry of the neuron in the vicinity of the soma may be obtained from the study of the relative size of the A spikes originated in different branches. These observations, together with the presence of different sizes of A spikes, produced by orthodromic stimulation, provide evidence that spikes initiated at separate axonal "trigger zones" of Aplysia neurons may be conducted selectively to the effectors or other neurons innervated by the particular branch.     

Extra spike formation in sensory neurons and the disruption of afferent spike patterning

The peculiar pseudounipolar geometry of primary sensory neurons can lead to ectopic generation of "extra spikes" in the region of the dorsal root ganglion potentially disrupting the fidelity of afferent signaling. We have used an explicit model of myelinated vertebrate sensory neurons to investigate the location and mechanism of extra spike formation, and its consequences for distortion of afferent impulse patterning. Extra spikes originate in the initial segment axon under conditions in which the soma spike becomes delayed and broadened. The broadened soma spike then re-excites membrane it has just passed over, initiating an extra spike which propagates outwards into the main conducting axon. Extra spike formation depends on cell geometry, electrical excitability, and the recent history of impulse activity. Extra spikes add to the impulse barrage traveling toward the spinal cord, but they also travel antidromically in the peripheral nerve colliding with and occluding normal orthodromic spikes. As a result there is no net increase in afferent spike number. However, extra spikes render firing more staccato by increasing the number of short and long interspike intervals in the train at the expense of intermediate intervals. There may also be more complex changes in the pattern of afferent spike trains, and hence in afferent signaling.     

Long-range afferents in the rat spinal cord. 1. Numbers, distances and conduction velocities.

The caudal extent of the penetration of primary afferent axons from the T12 and L1 dorsal roots and sural nerve has been investigated in adult decerebrate spinal rats. Microelectrode stimulation at the root entry zone (REZ) and at further caudal points in the spinal cord was used to generate antidromic action potentials in single fibres recorded in dorsal roots or peripheral nerves. A total of 209 units were recorded in T12 and L1 dorsal roots and 27% of these could be antidromically activated 10 mm caudal to the REZ. Fifteen percent of the units could be stimulated at the L4-5 border, 15 mm caudal to the T12 segment whereas 4.5% of the axons could be stimulated 25 mm caudally in the S4 segment, 11 segments caudal to the entry segment. Similar recordings made from units in the sural nerve showed that of all the sural axons that penetrated to the L6 segment 50%, 18% and 2% of these reached the S1, S2 and S4 segments respectively. The conduction velocities of these units were clearly in the A-beta range when recorded in the nerve but decreased on entering the spinal cord and were reduced by 83% at their caudal end point. The results show that substantial numbers of primary afferents have long-ranging caudal branches in areas beyond the regions of known postsynaptic effects. The functions of these caudal projections are unclear but they may represent a potential substrate for the development of functional connections under conditions of disease or denervation.     

Evoked Activity of Afferent and Efferent Fibers of the Sciatic Nerve in Rats under Conditions of Experimental Hyperthyroidism

In Wistar albino rats with experimental hyperthyroidism (HTh) and control animals, we measured parameters of the responses evoked in peripheral segments of the ventral and dorsal roots (VR and DR, respectively) by stimulation of the sciatic nerve. We found that the chronaxia of the afferent fibers of the sciatic nerve in HTh animals is shorter, while the duration of the mass action potential (AP) in the DR is somewhat longer than in the control. Under conditions of HTh, the excitation threshold of the efferent fibers became higher, the chronaxia decreased, and the second high-amplitude component could appear in the AP recorded from the VR. Possible mechanisms of changes in the excitability of afferent and efferent fibers of the sciatic nerve and specific features of the AP recorded from the VR under HTh conditions are discussed. In particular, we consider the possibility of ephaptic spreading of excitation in VR fibers under HTh conditions.     

Electrical excitability of the soma of sensory neurons is required for spike invasion of the soma,but not for through-conduction

The cell soma of primary sensory neurons is electrically excitable, and is invaded by action potentials as they pass from the peripheral nerve, past the dorsal root ganglion (DRG) and toward the spinal cord. However, there are virtually no synapses in the DRG, and no signal processing is known to occur there. Why, then, are DRG cell somata excitable? We have constructed and validated an explicit model of the primary sensory neuron and used it to explore the role of electrical excitability of the cell soma in afferent signaling. Reduction and even elimination of soma excitability proved to have no detectable effect on the reliability of spike conduction past the DRG and into the spinal cord. Through-conduction is affected, however, by major changes in neuronal geometry in the region of the t-junction. In contrast to through-conduction, excitability of the soma and initial segment is essential for the invasion of afferent spikes into the cell soma. This implies that soma invasion has a previously unrecognized role in the physiology of afferent neurons, perhaps in the realm of metabolic coupling of the biosynthesis of signaling molecules required at the axon ends to functional demand, or in cell-cell interaction within sensory ganglia. Spike invasion of the soma in central nervous system neurons may play similar roles.     

Target cell contact suppresses neurite outgrowth from soma–soma paired Lymnaea neurons

Neurite extension from developing and/or regenerating neurons is terminated on contact with their specific synaptic partner cells. However, a direct relationship between the effects of target cell contact on neurite outgrowth suppression and synapse formation has not yet been demonstrated. To determine whether physical/synaptic contacts affect neurite extension from cultured cells, we utilized soma–soma synapses between the identified Lymnaea neurons. A presynaptic cell (right pedal dorsal 1, RPeD1) was paired either with its postsynaptic partner cells (visceral dorsal 4, VD4, and Visceral dorsal 2, VD2) or with a non‐target cell (visceral dorsal 1, VD1), and the interactions between their neurite outgrowth patterns and synapse formation were examined. Specifically, when cultured in brain conditioned medium (CM, contains growth‐promoting factors), RPeD1, VD4, and VD2 exhibited robust neurite outgrowth within 12–24 h of their isolation. Synapses, similar to those seen in vivo, developed between the neurites of these cells. RPeD1 did not, however, synapse with its non–target cell VD1, despite extensive neuritic overlap between the cells. When placed in a soma–soma configuration (somata juxtaposed against each other), appropriate synapses developed between the somata of RPeD1 and VD4 (inhibitory) and between RPeD1 and VD2 (excitatory). Interestingly, pairing RPeD1 with either of its synaptic partner (VD4 or VD2) resulted in a complete suppression of neurite outgrowth from both pre‐ and postsynaptic neurons, even though the cells were cultured in CM. A single cell in the same dish, however, extended elaborate neurites. Similarly, a postsynaptic cell (VD4) contact suppressed the rate of neurite extension from a previously sprouted RPeD1. This suppression of the presynaptic growth cone motility was also target cell contact specific. The neurite suppression from soma–soma paired cells was transient, and neuronal sprouting began after a delay of 48–72 h. In contrast, when paired with VD1, both RPeD1 and this non‐target cell exhibited robust neurite outgrowth. We demonstrate that this neurite suppression from soma–soma paired cells was target cell contact/synapse specific and Ca²⁺ dependent. Specifically, soma–soma pairing in CM containing either lower external Ca²⁺ concentration (50% of its control level) or Cd²⁺ resulted in robust neurite outgrowth from both cells; however, the incidence of synapse formation between the paired cells was significantly reduced. Taken together, our data show that contact (physical and/or synaptic) between synaptic partners strongly influence neurite outgrowth patterns of both pre‐ and postsynaptic neurons in a time‐dependent and cell‐specific manner. Moreover, our data also suggest that neurite outgrowth and synapse formation are differentially regulated by external Ca²⁺ concentration. © 2000 John Wiley & Sons, Inc. J Neurobiol 42: 357–369, 2000     

Commissural neurons of the electrosensory lateral line lobe of Apteronotus leptorhynchus: morphological and physiological characteristics

Extracellular injections of horseradish peroxidase were used to label commissural cells connecting the electrosensory lateral line lobes of the weakly electric fish Apteronotus leptorhynchus. Multiple commissural pathways exist; a caudal commissure is made up of ovoid cell axons, and polymorphic cells' axons project via a rostral commissure. Intracellular recording and labeling showed that ovoid cells discharge spontaneously at high rates, fire at preferred phases to the electric organ discharge, and respond to increased receptor afferent input with short latency partially adapting excitation. Ovoid cell axons ramify extensively in the rostro-caudal direction but are otherwise restricted to a single ELL subdivision. Polymorphic cells are also spontaneously active, but their firing is unrelated to the electric organ discharge waveform. They respond to increased receptor afferent activity with reduced firing frequency and response latency is long. Electrical stimulation of the commissural axons alters the behavior of pyramidal cells in the contralateral ELL. Basilar pyramidal cells are hyperpolarized and nonbasilar pyramidal cells are depolarized by this type of stimulation. The physiological results indicate that the ovoid cells participate in common mode rejection mechanisms and also suggest that the ELLs may function in a differential mode in which spatially restricted electrosensory stimuli can evoke heightened responses.Abbreviations ccELL caudal commissure of the ELL - CE contralaterally excited - DML dorsal molecular layer - ELL electrosensory lateral line lobe - EOD electric organ discharge - HRP horseradish peroxidase - IE ipsilaterally excited - MTI mouth-tail inverted - MTN mouth-tail normal - rcELL rostral commissure of the ELL - TRI transverse inverted - TRN transverse normal     

Effect of preliminary mechanical and antidromic influences on orthodromic activity in C fibers from mechanoreceptors of the cat skin

Analysis of afferent activity in unmyelinated fibers of a cutaneous nerve was carried out by the colliding impulses method in cats. The effect of antidromic excitation of the nerve and mechanical stimulation of the receptors on subsequent orthodromic activity during stretching of the skin was investigated. Both these factors were shown to reduce subsequent orthodromic activity evoked by testing stimulation. The reduction in activity was greatest 10–15 sec after stimulation. The duration of the inhibitory effect was greater after mechanical than after antidromic stimulation. Combined mechanical stimulation and antidromic excitation resulted in a greater decrease of afferent activity and an increase in the time of its recovery. An increase in the frequency of antidromic excitation potentiated the inhibitory effect of preliminary stimulation on orthodromic activity in C fibers.Research Institute of Applied Mathematics and Cybernetics, N. I. Lobachevskii Gor'kii State University. Translated from Neirofiziologiya, Vol. 9, No. 3, pp. 307–312, May–June, 1977.     

Centrifugal and centripetal connections of the cat visual cortex

In experiments on curarized cats unit responses in the dorsal lateral geniculate body to stimulation of various zones in area 17 of the visual cortex were analyzed. Of all cells tested 69% were found to respond antidromically and 8% orthodromically; in 7.6% of cells IPSPs occurred either after an initial antidromic spike or without it. The velocities of conduction of excitation along the corticopetal fibers of the optic radiation varied from 28 to 4.3 m/sec, but the three commonest groups of fibers had conduction velocities of 28–19, 14–12, and 10–9.5 m/sec. A difference between latent periods of antidromic responses of the same neurons was found to stimulation of different zones of the visual cortex; this indicates that axons of geniculo-cortical fibers split into several branches which form contacts with several neurons in area 17 of the visual cortex. The degree and possible mechanisms of cortical influences on neurons of the lateral geniculate body are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 8, No. 3, pp. 243–249, May–June, 1976.