Spike Responses Of Neurons Research Articles

Neurons of the cat sensorimotor cortex: Responses related to differentiation of monomodal and heteromodal conditioning signals

Spike responses of the sensorimotor cortex neurons were studied in chronic experiments on cats trained to perform an operant reflex, a placing movement triggered by application of the distant stimuli. The responses recorded under conditions of differentiation of sound tones of various frequencies and of a heteromodal complex (light + tone) from its components were compared. The responses recorded from 125 neurons in 3 animals were analyzed. No neurons selectively responding either to monomodal or to heteromodal signals were found. Forty-five cells responded to positive signals by excitation or inhibition, irrespective of the signal modality. The shortest latencies of these responses were 30 and 40 msec, respectively. When inhibitory stimuli were applied, these neurons either generated much weaker responses, or did not respond at all. A correlation was found between the level of response depression and the level of differentiation of the signals by an animal. These findings allow us to hypothesize that the sensorimotor cortex does participate in differentiation of sensory signals, providing preparation for switching on the motor response after a positive stimulus or suppression of such a response after a negative one.

Non-reciprocal cross-adaptation of spiking responses of individual olfactory receptor neurons of spiny lobsters: evidence for two excitatory transduction pathways

Single-unit spiking responses of 72 olfactory receptor neurons (ORNs) in the olfactory organ of the spiny lobster Panulirus argus were recorded extracellularly during presentation of a set of seven odorant stimuli (adenosine-5′-monophosphate, ammonium chloride, betaine, l-cysteine, l-glutamate, d,l-succinate and taurine) and analyzed in order to evaluate the response specificities of single ORNs and the independence of receptor sites. Individual ORNs often had narrow excitatory response spectra, but the most excitatory compound was different from neuron to neuron. These results suggest that these compounds can exert most of their excitatory effects through relatively independent receptor site types. To determine the relative independence of excitatory transduction processes in single ORNs for these stimuli, single-unit spiking responses of these neurons under conditions of self- and cross-adaptation were analyzed. The results demonstrate extensive cross-adaptation between pairs of the seven stimuli. When averaged across all neurons and all cross-adaptation conditions, cross-adaptation resulted in a mean reduction of 81% of the unadapted response. However, there were differences in the degree and pattern of adaptation for different pairs of compounds and for different neuron types (defined by most excitatory or ‘best’ chemical). For a given neuron type, there were significant levels of non-reciprocal cross-adaptation: neurons cross-adapted more when adapted to their best chemical than when adapted to their non-best chemicals. These results suggest the existence of two excitatory transduction pathways within an olfactory receptor neuron: one pathway activated exclusively by the best chemical and a second pathway activated by a broader spectrum of chemicals.

Effect of response inhibition in cat motor cortex neurons during the expectation of conditioning stimulus

The spike responses of the motor cortex neurons (area 4) associated with forelimb movement were studied in awake cats earlier trained to perform placing motor reactions. Responses produced by the same neurons were compared in two situations: 1) when a sound-click conditioning stimulus (CS) was applied in isolation; 2) when a CS followed a preliminary warning stimulus (WS), a light flash, with a 100–1000 msec delay. During the reflex initiation by combined action of the WS and CS, response components that occurred prior to the placing movement (PM) performance under isolated CS action weakened and arrived 50–150 msec later; yet, response components that appeared in the same situation simultaneously with PM onset or later remained unchanged. PM latent periods were not changed when WS was applied. The temporal interval between WS and CS was characterized by depression of neuronal activity; depression duration was determined by the interstimulus delay. It is conceivable that the described transformations in spike responses of cortical neurons occurred due to changes in the sensory direction of the animal's attention; this direction, in all cases, is a crucial factor in the formation of neuronal activity in the cortex.

Neuronal spike response produced in the feline thalamic reticular nucleus by instrumental conditioned reflex

Spike response was investigated in 156 units of the thalamic reticular nucleus (RN) during performance of the instrumental feeding reflex of lever-pressing. This response consisted of lead and lag phases. Latency of the lead phase of response varied between 10 and 100 msec and total duration of response between 50 and 250 msec; minimum latency of the lag phase: 100–300 msec. Initial response to a conditioning clicking sound was found in 27 units, of which 26 showed excitation and the remaining single unit an inhibitory-excitatory pattern. The lag stage of response associated with performance of conditioned lever-pressing was found in 134 neurons, of which 115 showed an excitatory pattern, 19 displayed inhibition and the remaining 22 units failed to respond. The lag phase of response preceded the onset of conditioned reflex movement (CRM) in 30 neurons. A total of 118 neurons responded between the onset of CRM and the point of lever-pressing. It was concluded that the RN plays a part in perception of the conditioned signal as well as producing and controlling performance of CRM.

Inhibitory effects of external stimulation on neuronal spike response in the cat association cortex during conditioning

The effects of external stimulus presentation on neuronal spike response induced in area 7 by conditioned and unconditioned stimuli were investigated in waking cats. Pointing reaction arising in response to the action of external stimuli led to the phasing out of spike response to unconditioned cutaneous stimulation, as well as depression or complete suppression of response in area 7 neurons to conditioned stimulus presentation, despite occurrence of conditioned reflex motion. External stimuli no longer exerted an inhibitory effect on the neuronal spike response induced by protracted use of conditioned and unconditioned stimuli. The fact that the conditioned lever-pressing reflex could occur either following or without neuronal spike response in area 7 would raise the question of whether neurons belonging to this area always participate directly in the triggering and occurrence of conditioned reflex motion.

Changes produced in neuronal excitability by subthreshold depolarization as a possible mechanism of interval selective relationships within the central nervous system

A neuronal process was identified inLymnaea stagnalis nerve cells which may be viewed as one of the mechanisms underlying the interval selectivity previously described in research into the functional relationships between mammalian brain cells. This process takes the form of regularly-occurring changes in excitability resulting in a high probability (of 0.6–1) of neuronal spike response to what had previously been subthreshold depolarizing current pulses following similar subthreshold (conditioning) pulses at intervals specific to each individual neuron. It was found that the cycle of change in neuronal excitability following threshold depolarization did not arise from temporal summation of electrotonic local or postsynaptic neuronal potentials; it was an endogenous (cytoplasmic) process insensitive to transmitter (acetylcholine) application but altering irreversibly under the effects of bombesin, one of the modulator peptides.

Long-term posttetanic changes in neuronal spike responses of the cat sensomotor cortex.

Long-term posttetanic changes in neuronal spike responses of the cat sensomotor cortex.

Long-term posttetanic changes in neuronal spike responses of the cat visual cortex.

The phenomenon of long-term posttetanic potentiation has now been found in several different structures of the CNS [1-5] and it has been studied most fully in hippocampal neurons. The present writers have shown that monosynaptic responses of neurons of the cat motor cortex also undergo long-lasting posttetanic changes in the form of potentiation [i] or depression. This paper gives data on long-term posttetanic changes in neuronal spike responses in the cat visual cortex.

Spread of excitation in the upper layers (I–III) in an auditory cortex slab isolated from the cat

Research was performed on an auditory cortex slab isolated from unanesthetized immobilized cats after severing of the lower layers, preserving intact a “bridge” one, two, or three upper cortical layers under the pial membrane; the spike response of neurons on this slab to stimuli applied at the other side of this additional incision was observed. It was found that stimulation applied to level IV produces an excitatory wave which spreads to the upper cortical layers, leading to mono- and polysynaptic activation of neurons at all levels via the apical dendrite of pyramidal cells. Antidromic activation of layer IV neurons together with an especially high proportion of monosynaptic response was observed with the three upper cortical layers remaining intact. A possibly crucial role in the horizontal spread of excitation within the cerebral cortex of a major bundle of axons from cortical interneurons is discussed in this article.

Influence of food motivation on neuronal response of the cat somatic cortex during instrumental reflex

Spike activity in neurons of areas 3 and 4 was investigated in experiments on cats during the conditioned reflex response of placing the paw on a support both before and after feeding ad libitum. Ingestion of a feed consisting of a rapidly absorbed glucose dairy mix did not prevent the reflex taking place if the animals' favorite food was used as reinforcement. Background activity increased in two-thirds of the neurons after the feed; the tonic constituent of neuronal response declined substantially and repeated contraction of the biceps occurring at the same rate as locomotor movements disappeared. Difference in latency of response produced by the conditioned stimulus in the same neurons before and after feeding measured 50–300 msec during the experiment. Measurements of latency of placing motion remained largely unchanged. Changes in the latency of neuronal spike response were thus found to be interrelated with the intensity of the animal's motivational excitation. It is suggested that fluctuations in degree of food motivation lead to changes in cortico-subcortical relationships responsible for initiation and performance of conditioned movements in these animals.

Calcium-induced long-term potentiation in the hippocampal slice: Characterization of the time course and conditions

A transient increase in extracellular calcium concentration causes a long-lasting enhancement of radiatum fibers evoked excitatory postsynaptic potential and population spike responses of CA1 pyramidal neurons which resembles long-term potentiation (LTP). The duration of this potentiation is much longer than described previously and is probably limited by the survival of the preparation itself (>8 hr). Therefore, Ca-induced LTP can be used for the investigation of a postulated late phase of LTP. Ca effects were activity-independent, since the subsequently evoked responses were facilitated even when the presynaptic fibers were not concurrently stimulated during or immediately after superfusion with the high Ca medium. In contrast, if too frequent testing of the synaptic input was done during the high Ca pulse, a short lasting depression instead of potentiation was observed. A lower extracellular magnesium concentration in the standard medium (1.3 instead of 2.0 mM MgSO 4) prevents the potentiation of the EPSP at least for the first few hours. Presumably, both tetanus- and Ca-induced LTP share some common mechanisms, since an additional tetanization after Ca induction was not followed by an additional LTP. Compared to the potentiation following tetanization, the Ca-induced LTP was, however, not accompanied by a potentiation of the EPSP/spike ratio within the range of the population spike threshold intensity.

Involvement of mesencephalic reticular formation neurons in cat conditioned reflexes

Traditional defensive and operant food reflexes were used to investigate neuronal responses of the mesencephalic reticular formation. It was found that these neurons may be divided into different groups according to function, depending on how they respond to positive conditioning stimuli. Of the two main groups of neurons with sustained tonic reactions one is activated in response to positive acoustic conditioning stimulation; it no longer reacts to the same stimulus after extinction of the reflex, while the other only becomes involved in response to positive stimulation accompanying the initiation of movement. Neurons belonging to the second group begin to respond directly to acoustic stimulation after extinction of the conditioned reflex. Neurons of the mesencephalic reticular formation can thus exercise additional tonic ascending effects both in the production and inner inhibition of the conditioned reflex. The group of neurons with a phasic reaction, i.e., a double response (a direct response to sound and another produced by movement) displayed a drop in spontaneous activity during the shaping of inhibition of differentiation and of extinction in particular. It was found that the initial changes in the spike response of reticular formation neurons during conditioning and pseudo-conditioning are similar. There are thus grounds for stating that neurons of the mesencephalic reticular formation participate in the shaping, production, and inner inhibition of traditional and operant conditioned reflexes in a differentiated capacity rather than as a population reacting identically.

Model investigation of neuronal mechanisms of discrimination between two tactile stimuli in snails

Synaptic and spike responses of neurons were studied in a two-layered model based on a study of the structure of the neuronal net of the defensive reflex in snails [4], using a technique of simultaneous application of two equal stimuli. When the points of application of the two stimuli were brought closer together the bimodal distribution of synaptic and spike responses of the efferent neurons of the model gradually changed into unimodal. The character of quantitative changes in synaptic and spike responses of neurons of the model, as the points of application of the two stimuli move closer together, reflects well the character of changes in effector responses of the snail's foot under experimental conditions as the distance between the two simultaneously acting tactile stimuli is reduced. After removal of the CNS, responses of contraction of the foot muscles of the snail become less accurate (they have a more diffuse maximum). It is suggested that the difference between functions of the central and peripheral nerve nets in molluscs lies in differences in the degree of accuracy of performance of the reflex response.

Neuronal responses in area 7 of the cat cortex during defensive conditioning

Responses of neurons in area 7 of the parietal association cortex during and after formation of a defensive conditioned reflex to sound were recorded in waking cats. Changes in spike responses of the neurons as a result of the onset of conditioned reflex limb movements were observed in 68% of neurons. Spike responses of neurons formed as a result of learning appeared only if conditioned-reflex limb movements appeared, and they were not observed if, for some reason or other, movements were absent after presentation of the positive conditioned stimulus or on extinction of the reflex. Responses of 46% neurons to conditioned stimulation preceded the conditioned-reflex motor responses by 50–450 msec. The remaining responding neurons were recruited into the response after the beginning of movement. Characteristic spike responses of neurons to the conditioned stimulus appeared 500–900 msec before the beginning of movement and, in the case of appearance of special, "prolonged" motor responses of limb withdrawal, evoked by subsequent reinforcing stimulation.

Receptive field dynamics of neurons in the cat visual cortex and lateral geniculate body

Spike responses of single neurons in the primary visual cortex and lateral geniculate body to random presentation of local photic stimuli in different parts of the receptive field of the cell were studied in acute experiments on curarized cats. Series of maps of receptive fields with time interval of 20 msec obtained by computer enabled the dynamics of the excitatory and inhibitory zones of the field to be assessed during development of on- and off-responses to flashes. Receptive fields of all cortical and lateral geniculate body neurons tested were found to undergo regular dynamic reorganization both after the beginning and after the end of action of the photic stimulus. During the latent period of the response no receptive field was found in the part of the visual field tested, but later a small zone of weak responses appeared only in the center of the field. Gradually (most commonly toward 60–100 msec after application of the stimulus) the zone of the responses widened to its limit, after which the recorded field began to shrink, ending with complete disappearance or disintegration into separate fragments. If two bursts of spikes were generated in response to stimulation, during the second burst the receptive field of the neuron changed in the same way. The effects described were clearly exhibited if the level of background illumination, the intensity of the test bars, their contrast with the background, duration, angles subtended, and orientation were varied, although the rate and degree of reorganization of the receptive field in this case changed significantly. The functional importance of the effect for coding of information about the features of a signal by visual cortical neurons is discussed.

Comparative analysis of the role of the second and first somatosensory cortical areas in somatic responses of the medullary reticular formation in rats

Evoked potentials and unit activity in the medullary reticular formation were investigated in unanesthetized, curarized rats during cold blocking or after extirpation of the cortical representation of one of the stimulated limbs. Local cooling or extirpation in area SII, unlike blocking of area SI, leads to a small (up to 30%) decrease in amplitude and a very small change (up to 10 msec) in the temporal parameters of evoked potentials arising in the reticular formation in response to electrodermal stimulation of the contralateral limb, whose representation in the cortex was blocked. Predominance of corticofugal influences from SI over those from SII was discovered both in experiments with evoked potentials and during analysis of somatic spike responses of reticular formation neurons. Corticofugal control over activity of the medullary reticular formation in rats exerted by neuronal mechanisms of somatosensory areas SII and SI thus differs both qualitatively and quantitatively.

The action of piracetam on the electrical activity of the hippocampal slice preparation: A field potential analysis

The action of various doses of piracetam on the electrical responses of in vitro hippocampal slices from the rat was investigated. Piracetam increased dose dependently the amplitude of the population spike response of pyramidal neurons evoked by stimulation of the stratum radiatum. Piracetam began to affect the population spike at a dose of 100 μM and consistently increased it at a dose of 1 mM. The drug had a rapid onset of action and recovery was seen within a few minutes following its removal. The response of the pyramidal neurons to antidromic stimulation was not affected by piracetam at concentrations up to 50 mM. Also, neither the amplitude nor the slope of the dendritic response was changed by the drug. Piracetam also did not affect either posttetanic or long-term potentiation of synaptic potentials. These findings suggest that the drug does not act through depolarization of the pyramidal cells or potentiation of the synaptic processes located on their dendrites. The action of piracetam on the feedforward- and feedback-mediated inhibition of pyramidal cells by basket cells was investigated in several series of experiments. At concentrations of 1 and 10 mM, piracetam did not change either of the two types of inhibitory mechanisms. Several hypothetical sites of action of piracetam are discussed.

Calcium-induced long-term potentiation in the hippocampus

The effect of a transient increase in extracellular calcium concentration on the Schaffer collateral-commissural evoked excitatory postsynaptic potential and population spike responses of CA1 pyramidal neurons was investigated using the rat in vitro hippocampal slice preparation. Brief exposure of slices (5–10 min) to twice the normal concentration of calcium (4 mM) induced a marked potentiation of both the excitatory postsynaptic potential and population spike that could persist for at least 3 h. No long-term changes were observed in either the presynaptic fiber volley or antidromically evoked CA1 population spike, indicating that the potentiation could not be attributed to an increase in the number of fibers activated or a generalized increase in cellular excitability. The response of CA1 pyramidal neurons to the iontophoretic application of l-glutamate in the apical dendritic zone was also unaffected after exposure to high calcium perfusate, suggesting a lack of alteration in membrane excitability or receptor sensitivity restricted to the region of synaptic input. In addition, total intracellular calcium content of individual slices, measured by atomic absorption spectrophotometry, was significantly increased for at least 1 h following return to the control medium. These data indicate that brief exposure of in vitro hippocampal slices to a high extracellular calcium concentration results in a long-term increase in synaptic efficacy which is similar in many respects to long-term potentiation induced by tetanic stimulation of hippocampal excitatory afferents. The results further suggest that the mechanisms underlying calcium-induced long-term potentiation may reside in presynaptic components and involve an enhanced transmitter release.

Selective effects of enkephalin on electrical activity of the in Vitro hippocampal slice

The effect of infusion of small concentrations of d-Ala-d-Leu-enkephalin on electrical responses to stimulation of the in vitro hippocampal slice preparation was investigated. Concentrations of enkephalin as low as 10 nM dramatically increased the magnitude of the population spike response of pyramidal cell neurons to stimulation of their Schaffer-commissural afferents. This effect had a rapid onset and was quickly eliminated when the peptide was removed from the perfusion medium. Enhancement of the population spike was blocked by the opiate receptor antagonist naloxone, which, by itself, did not affect either synaptic potentials or the population response. In contrast even relatively high concentrations of the enkephalin produced no detectable effect on either the synaptic potentials recorded in the dendrites or the antidromic response of the pyramidal cells to stimulation of their axons. These findings suggest that enkephalin attenuates the effect of either a feedback or feed-forward inhibitory system which is usually activated by stimulation of the Schaffer-commissural pathways.

Effect of stimulus intensity on spike responses of crayfish receptor neurons

The character of spike responses of receptor neurons of n. telsoni ventralis to changes in the direction, velocity, and angle of displacement of the uropod hairs was studied in experiments onAstacus leptodactylus. For all receptor neurons the latent period is inversely proportional and the mean impulse frequency directly proportional to the velocity of hair displacement. Selective sensitivity of different receptor neurons to the direction of motion and also to a change in the angle of displacement of the hair was discovered; certain zones of angular displacements of the hair characterized by a maximal response of the receptor neuron were distinguished.