Role of the parietal associative cortex in "counting" behavior of dogs

Influence of the combined and isolated lesions of areas 5 and 7 of the parietal cortex on the counting behavior was studied in experiments with 6 dogs. Instrumental feeding reaction (lifting and placing the forepaw on the foodwell) was established. The positive conditioned stimulus was a series of 5 clicks with variable interclicks intervals and the negative (non-reinforced) conditioned stimulus was a series of 3 clicks, so that asymmetrical differentiation was elaborated. Combined bilateral lesions of areas 5 and 7 and an isolated lesion of area 5 resulted in a severe impairment of the numerical discrimination for two months, whereas the isolated lesion of area 7 did not lead to any problems in differentiation. The conclusion was made that area 5 is critical for numerical discrimination of sequential stimuli.     

Temporal pattern sensitive and nonsensitive responses in the cat's retinal ganglion cells

In order to characterize temporal pattern sensitivity in the cat ganglion cells, a new analysis technique by semi-Markov models which was developed in the previous papers (Tsukada et al., 1975–1977) was applied to input-output relations of the receptive-field. Three types of statistical spot stimuli positioned in the center region of receptive fields were used. Each type of stimulus has an identical histogram in the inter-stimulus intervals and therefore the same mean and variance, but different correlations between adjacent inter-stimulus intervals (Type 1, positive; Type 2, negative; and Type 3, independent processes). From the output spike trains of cat retinal ganglion cells to each stimulus, mean, variance, and histogram were computed. As the result of investigating these data, we could draw the following conclusion from the resultant output interval histograms. The receptive-field-center responses of cat ganglion cells can be classified into two groups (Types L and N) according to the difference of responsiveness to the three types of statistical spot stimuli. A Type L response has the same histogram in interspike intervals for all three stimuli, and is not sensitive to the temporal pattern, while a Type N response has three different forms depending on each type of stimulus showing high sensitivity to the temporal pattern. These results were also simulated by the Markov chain model and discussed with relation to neural coding and classification of ganglion cell types.     

Reversal of signal significance of stimuli in the avoidance reflex]

The course of reversal learning in go -- no go avoidance reflex differentiation in both cats and dogs was analysed in terms of interrelations between drive and instrumental reflex activity. When signalling properties of the conditioned stimuli were reversed, marked changes in the drive state of experimental animals occurred. The increase of the fear drive influenced both the transfer of the instrumental response to the new positive conditioned stimulus and the extinction of the response to the previously positive stimulus. The quality of the conditioned stimuli and their reflexogenic strength exerted clear effects on the fear drive and on the course of reversal learning. Prefrontal lesions affected the drive state and the conditioned reflex activity of the animals during the reversal learning.     

Excitability cycles of direct cortical responses during elaboration of conditioned defense reflexes in dogs]

Direct cortical responses (DCRs) to paired stimuli were studied in chronic experiments in dogs during elaboration of classical and instrumental defensive conditioned reflexes. The DCRs were recorded with 20 to 250 ms intervals between stimuli. Paired and single electrical stimulations of the middle suprasylvian gyrus given with a frequency of one per second were used as conditioned stimuli and were reinforced in a similar way. During electrical cutaneous stimulation of the dog's paw and to an even greater extent during isolated action of the conditioned stimulus the initial negativity of the testing DCR became shorter and the degree of its depression diminished. In the case of a following period of facilitation, its degree became greater. It was higher at a distance of 4 to 5 mm from the point of stimulation than at a distance of 2 to 3 mm. During isolated action of the conditioned stimulus, the degree of facilitation was higher than at the period of the possible action of the unconditioned stimulus. The greatest shorterning of the DCR excitability cycle was observed immediately before and during the conditioned lifting of the dog's paw. Excitability cycles of DCR, and possibly of other evoked potentials as well, are a more sensitive indicator of the function state of the cerebral cortex than responses to single stimuli. For this reason it appears promising to use them in studying conditioned reflexes.     

Neuronal correlates of the activation of the brain-reinforcing system in dogs

In analysis of neuronal activity of g. proreus of dogs brain in response to positive and differentiation conditioned cutaneous stimuli, in 44% of 78 studied neurons "late responses" appeared in the form of impulse activity intensification elicited by a positive conditioned stimulus after the end of eating and by a differentiation one--during conditioned signal action; it persisted in different neurones for 30--60 s. By means of various functional tests it has been shown that these reactions are a neurophysiological correlate of animals emotional state gradient.     

Vector representation of associative learning

I. P. Pavlov [12] has shown that conditioned reflexes are selective both with respect to conditioned stimuli and to conditioned reflexes elicited by those conditioned stimuli. At the neuronal level selective aspects of conditioned stimuli are based on detectors selectively tuned to respective stimuli. The selective aspects of conditioned reflexes are due to command neurons representing specific unconditioned reflexes. It can be assumed that conditioned reflexes result from association between selective detectors and specific command neurons. The detectors activated by a conditioned stimulus constitute a combination of excitations--a detector excitation vector. The detector excitation vector acts on a command neuron via a set of plastic synapses--a synaptic weight vector. Plastic synapses are modified in the process of learning making command neuron selectively tuned to a specific conditioned stimulus. The selective tuning of a particular command neuron to a specific excitation vector referred to a conditioned stimulus is a basis of associative learning. The probabilities of conditioned reflexes elicited by conditioned and differential stimuli implicitly contain information concerning excitation vectors that encode respective stimuli. Contribution of the vector code to associative learning was explored combining differential color conditioning with intracellular recording from color-coding neurons. It was shown that colors in carps and monkeys are represented on a hypersphere in the four-dimensional space similar to human color space. The basis of the color space is constituted by red-green, blue-yellow, brightness and darkness neurons.     

Microelectrode investigations of learning phenomena in snail (Helix pomatia) neurones.

We have examined changes of postsynaptic potentials and of pattern activity of the identified silent and oscillatory snail neurones in Helix pomatia during conditioning. Local changes of EPSP or IPSP have been recorded during association following the first stimulus in the silent cells, whereas spike discharges could be observed in response to the 2nd stimulus. In the oscillatory neurones changes of pattern activity have been recorded following the 2nd stimulus, while the first stimulus proved to be ineffective. The formation of temporary connections of snail neurones seemed to be a specific phenomenon, because it was necessary to pair stimuli of different inputs for the development of these modifications. These plastic changes seemed to depend on the interstimulus as well as on the intertrial intervals. Our experimental data underline the probable role of the stimulus parameters and of the electrical properties of neurones during the formation of learned neuronal responses.     

Classical heart-rate conditioning and differentiation of visual CS with an appetitive UCS in pigeons

A tachycardic response was classically conditioned in thirsty pigeons using water as an unconditioned stimulus and localized lights as conditioned stimuli. This was successful in 6 out of 9 birds. In one experiment the subjects had their heads immobilized so that the positions of the stimuli within the visual field could be precisely specified. When the stimuli were shifted from the binocular to the monocular field and vice versa there was no initial transfer of a conditioned differentiation between a white and a green light.     

The dependence of the neuronal reactions of the sensorimotor cortex to a simultaneous complex stimulus on the level of the differentiation of its components]

On alert animals the change was studied of the neuronal activity of the sensorimotor cortical area of cats brain in dependence on the level of differentiation of the components of simultaneous heteromodal complex stimulus. According to the character of this dependence and a number of other parameters two groups of neurones were singled out in the sensorimotor cortex. It was shown that parameters of reactions of all recorded neurones of the sensorimotor cortex to the positive conditioned signal were the first established after consolidation of the animal conditioned motor activity. In the course of elaboration following parameters changed: expressiveness, intensity, duration and value of latency. Reactions of neurones of both groups to the inhibitory stimuli were stabilized only after consolidation of the habit of differentiation. Responses of the first group neurones changed only by the pattern of discharge, while the responses of the second group neurones could change by expressiveness of response, its sign, duration and value of latency. Oscillations of the differentiation level after finishing of the elaboration of inhibitory conditioned reactions affected only the responses of the second group neurones to complex components.     

Effect of sectioning the brachia of the inferior colliculi on the formation of auditory conditioned reflexes to amplitude-modulated stimuli

Elaboration of differentiation of tonal and amplitude-modulated stimuli with a frequency modulation of 5 Hz conducted on laboratory rats in five months after section of brachii colliculi proved to be impossible. The rate of correct reactions did not differ from the probability of random choice during 500 presentations of conditioned stimuli but significantly differed from the values obtained during differentiation of these signals in rats after control operation. Electrophysiological control showed that while in the control group and in intact animals at the presentation of amplitude-modulated stimuli with modulation frequency of 1-20 Hz summary potentials were recorded in the auditory cortex synchronized with their modulation frequency,--in rats after section of brachii colliculi only on- and off-responses were observed. It may be suggested that sensory providing of conditioned activity during the action of amplitude-modulated stimuli with low modulation frequency may be achieved only with the participation of specific pathways.     

Changes in cortical unit activity during selective reinforcement of a chosen interspike interval range

In experiments on immobilized unanesthetized rabbits selective automatic reinforcement of certain ranges of short and long interspike intervals in spontaneous unit activity recorded from single cortical units was given by means of a nociceptive stimulus. Analysis of postinterval histograms showed that the total number of intervals reinforceable by nociceptive stimuli is reduced as a result of the consequent reorganization of the firing pattern. If short intervals are reinforced the mean firing rate is reduced and the probability of appearance of long intervals immediately after the short reinforcements is increased. After reinforcement of long intervals the mean firing rate was increased in one group of cortical neurons, with a corresponding decrease in the total number of long intervals. In the other group of cells a decrease in the number of long intervals was accompanied by a decrease in the mean firing rate of the cells. It is postulated that reorganization of the firing pattern is determined by the level of synchronization of ascending activating influences and by their summation with phases of excitation and inhibition in reciprocally interacting neuron populations.Institute of Experimental Medicine, Academy of Medical Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 4, No. 4, pp. 339–348, July–August, 1972.     

The effect of an extra-stimulus on successive inhibition following differentiations]

The influence of a foreign stimulus on the aftereffect of inhibitory stimuli was studied in experiments on dogs with salivary conditioned reflexes. The dynamics of temperature changes in the parotid salivary gland was used as an indicator since the action of a differentiation stimulus was accompanied by a drop in its temperature; each differentiation, whether rough, medium or fine--had its own limit of the gland's temperature drop during intensification of differentiation inhibition. It has been found that the dynamics of the gland temperature changes in the trace pauses and in response to a foreign stimulus depends on the modality of the stimulus, the duration of its action in the experiment, the typological properties of the animal's nervous system and the fineness of differentiation.     

Statistical characteristics of spontaneous and evoked unit activity in the cat superior colliculus

Spontaneous and evoked unit activity of the superior colliculus was studied in cats immobilized with gallamine or with the spinal cord divided (encephalé isolé). Statistical parameters of unit activity were calculated in light and darkness and in the presence of a moving photic stimulus. Spontaneously active neurons were divided into four main groups: 1) with a mean interspike interval of about 300 msec, coefficient of variation 1.9, and with two modes in the autocorrelation histogram; 2) with similar statistical parameters but distinguished by correlation with saccadic eye movements; 3) histogram of interspike intervals exponential in type, autocorrelation histogram flat, mean interval and standard deviation about 100 msec; 4) with symmetrical distribution of interspike intervals, mean interval about 50 msec, coefficient of variation 0.4–0.5. Spontaneous activity in light and darkness differed only in the neurons of the second group. Comparison of the statistical parameters of the spontaneous and evoked activity shows that encoding of the stimulus could be carried out by neurons of groups 3 and 4. In some cases the mode of the histogram of interspike intervals and the standard deviation changed significantly and activity appeared regularly in the autocorrelation histogram.     

Dynamics of elaboration of a conditioned reflex and differentiations in the snail

It is shown that dynamics of percentage of conditioned food refusals by snails Helix pomatia and Helix lucorum is satisfactorily approximated by regression equation of exponential type with included coefficient reflecting the animals state before the beginning of learning. The ability is established of the snails to practically completely differentiate various alimentary conditioned stimuli. The introduction of differentiation always improved the reflex consolidation. Conditioned reactions to differentiation stimulus appeared at the elaboration stage and were absent at the stage of conditioned reflex consolidation.     

Functional differentiation of the theta-rhythm of the hippocampus in the cat

On the model of discriminating alimentary conditioned reflex with spatial separation of the sources of conditioned (CR, light flashes) and unconditioned (feeding trough) stimuli the theta-rhythm of the dorsal hippocampus was evaluated in cats. Two types of the theta-rhythm were observed in the spectrum of the hippocampal electrical activity: low-amplitude, consisting of a slow growing theta-waves (type I) and high-amplitude consisting of rapidly increasing theta-waves (type II). The type I theta-rhythm is sensitive to noradrenaline and correlates with behavioural forms directed immediately to the realization of alimentary motivation, while the type II is sensitive to serotonin and correlates with behavioural forms directed to the source of the conditioned stimulus. Enhancement of the type II theta-rhythm takes place during a delay of the expected conditioned stimulus. It is suggested that type I theta-rhythm reflects a level of activity of the brain structures connected with unconditioned mechanisms, with realization of biological motivations, while type II theta-rhythm is connected with conditioned reinforcing stimuli; it correlates with various forms of conditioned orienting reflexes and reflects the work of the "nervous model of reinforcing stimulus".     

Regression analysis of non-stationary discharges in neurons; Adaptation in the electrosensory afferent of dogfish

Interspike interval histograms, as usually regarded for the estimation of statistical variabilities in neuronal spike trains, were applied to non-stationary dynamic responses of a PD receptor. Sliding mean values were introduced describing the average receptor response on defined, recurrent stimuli; mean spike frequencies and interspike intervals were computed a) for fixed sequential analysis periods (of e.g. 500 ms), b) for analysis periods shifted by every consecutive interspike interval (thus the number of spikes being constant), and c) by fitting the dynamic responses for suitable analytic functions (e.g. exponential functions). With these methods variabilities in the non-stationary neuronal impulse patterns were investigated for electrosensory PD afferents in Lorenzinian ampulla of dogfish (Scyliorhinus canicula) with electric stimuli up to 50 nA and defined temperatures between 7° C and 25° C. In this temperature range all investigated ampullae were spontaneously active, the irregularities in neuronal discharges and averaged spike frequencies depended strongly on temperature, the latter showing maxima between 13° C and 19° C. In preparations with small disturbances we generally found static interspike interval histograms following approximatively a Gaussian distribution. The same was true for the momentary spike frequency and its deviation during the dynamic response to given electrical stimuli. A suprathreshold rectangular current (e.g.-0.5 nA) led to a marked but transient synchronisation in spike generation; the higher the stimulus strength, the smaller the standard deviation (s.d.) from mean spike frequency in the beginning of the dynamic response; during adaptation the s.d. increased up to that of the static response frequency. Relating, however, s.d. for different currents, times, and temperatures to the corresponding mean spike frequency led to fairly constant coefficients of variation; s.d. was approximatively a linear function of the sliding mean value even in the dynamic response of the electroreceptor (scaling).Supported by the Deutsche Forschungsgemeinschaft (Br 310/11)     

Low-frequency neuronal oscillations as operant processing of integrative mechanisms of conditioning and plasticity

It was found that neuronal activity of the rabbit somatosensory cortex could be operantly conditioned according to the given algorithm of biofeedback control of nociceptive stimulation leading to rising or lowering of firing rate of multiunit activity, or to changes of range of units interspike intervals, or neuronal firing patterns for conditional stimulus, which become operants. Neuronal interactions of its population with feedback and systems of cardiovascular and pain regulation lead to appearance of wave patterns, combining elements to integrative activity by conditioning. Low-frequency oscillations of neuronal assemblies within range of 0.02-0.8 Hz act as operant processing in mechanisms of system plasticity and conditioning.     

Classical conditioning of ventilatory responses in humans

A classical conditioning experiment, in which an auditory stimulus was paired with a hypoxic stimulus, was carried out on 34 normal subjects assigned to two groups (experimental and control). Each subject took part in one session divided into two phases, acquisition and test. In the acquisition phase, eight hypoxic and eight auditory stimuli were paired in the experimental group and unpaired in the control group. In the test phase, which was identical for the two groups, the hypoxic stimuli were suppressed and three purely auditory stimuli were presented. Significant differences between the two groups in ventilatory response to these auditory stimuli provided evidence for conditioning. In the control group, no significant changes were elicited by the auditory stimuli, whereas a conditioned increase in total cycle duration was observed in the experimental group. The conditioned response closely resembled the first component of the hypoxic response. Analysis of the pattern of the conditioned response, along with postexperimental interviews, strongly suggests that this response was not mediated by volitional factors.     

Spatial interactions and stimulus-response relations of unit responses evoked by somato-sensory stimuli in the feline caudal medullary reticular formation

Responses of neurons in the bulbar reticular area to separate and simultaneous stimulation of the forelimbs were recorded extracellularly in chloralose-anaesthetized cats. On increasing the stimulus intensity the number of spikes per response increased while the initial latency and interspike intervals decreased in accordance with the functional property of the neuron. Responses evoked by simultaneous stimulation displayed more spikes and a shorter latency than those evoked by separate stimuli of corresponding intensities. The differences in the responses evoked simultaneously and the sums of responses evoked separately showed characteristic distributions as a function of the latter. Three types of distribution were distinguished. The results indicate that stimulus-response relations play a determining role in the mechanism of spatial integration.     

Stimulus representation in SOP: II. An application to inhibition of delay

The componential extension of SOP accounts for conditioned response (CR) timing in Pavlovian conditioning by assuming that learning accrues with relative independence to stimulus elements that are differentially occasioned during the duration of the conditioned stimulus (CS). SOP, using a competitive learning rule and the assumption that temporal learning emerges via resolution of what is equivalent to an "AX+BX-" discrimination, predicts a progressive increase in the latency of the CR over training, or what Pavlov refer to as "inhibition of delay." Other componential models, which use noncompetitive learning rules, do not predict inhibition of delay. Either type of model makes the prediction indicated, independently of the length of the CS-unconditioned stimulus (US) interval. We report two experiments that demonstrated inhibition of delay when rabbits were trained with relatively long, but not with short, CS-US intervals. To account for this divergence, we assumed that the SOP stimulus trace involves two kinds of elements, some with a temporally distributed pattern of activity over the duration of the CS duration, and some with a randomly distributed pattern. This stimulus representation, not only allows for inhibition of delay with long but not short CS-US intervals, but in combination with SOP's performance rule deduces CR's with "Weber variability."