Formation of "caudate spindles" in the rabbit sensomotor cortex during ontogeny

Electrical activity of the sensomotor and visual areas of the neocortex during stimulation of the caudate nucleus was recorded in young rabbits aged 3–60 days and in adults. Single stimulation of the caudate nucleus was found to cause the appearance of characteristic bursts of spindle-like rhythmic activity ("caudate spindles"), described previously in cats and monkeys, in the adult rabbit cortex. The latent period of the caudate spindle was about 200 msec, its duration 1–3 sec, and the frequency of its oscillations of the order of 12 Hz. Caudate spindles were most marked in the sensomotor cortex of the ipsilateral hemisphere. In rabbits under 10 days old caudate spindles were not found even if the intensity of stimulation was increased many times. Starting from the age of 15 days bursts of rhythmic activity resembling caudate spindles, but with lower frequency (about 8 Hz), longer latent period (up to 350 msec), and also with a higher threshold, appeared in the sensomotor cortex. The definitive type of caudate spindles was established toward the end of the first month of postnatal life, corresponding to the time of formation and complication of conditioned-reflex activity in developing animals.Brain Institute, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 17, No. 1, pp. 11–15, March, 1985.     

Postnatal changes in the dorsal root reflex in the isolated spinal cord of the hamster,Mesocricetus auratus

Spontaneous activity has been demonstrated in the lumbar dorsal roots of isolated spinal cord preparations taken from animals ranging in age from 2 to 65 days. Peaks of activity were recorded at 2 and 5 weeks of age, with mean firing frequencies of 33 Hz and 28 Hz respectively. The firing frequency in weeks 3 and 4 was lower (15 Hz) as was the frequency in cords taken from animals older than 6 weeks. The pattern of the spontaneous dorsal root activity changed during the first 5 weeks of life. In cords taken from animals less than 10 days old, the roots fired single action potentials, producing a single broad peak in Inter Spike Interval plots (ISI). Dorsal root recordings made from cords taken from animals in weeks 2 and 3 of life exhibited both single spikes and bursts of action potentials. By the end of the third week of life, individual spike activityhad declined and the bursts of action potentials characteristic of the adult pattern had become dominant, producing a bimodal ISI plot. Cross correlation analysis of dorsal root and dorsal horn activity in lumbar segments up to five segments apart, revealed an increasing degree of correlation developing over the first 4 weeks of postnatal life. Dorsal horn responses to dorsal root stimulation in cords taken from young animals were prolonged, lasting in excess of 250 msec. In the third week of life, the duration of the excitatory component of the response was reduced to approximately 50 msec by the development of an inhibitory phase.     

Alpha-Band Rhythms in Visual Task Performance: Phase-Locking by Rhythmic Sensory Stimulation

Oscillations are an important aspect of neuronal activity. Interestingly, oscillatory patterns are also observed in behaviour, such as in visual performance measures after the presentation of a brief sensory event in the visual or another modality. These oscillations in visual performance cycle at the typical frequencies of brain rhythms, suggesting that perception may be closely linked to brain oscillations. We here investigated this link for a prominent rhythm of the visual system (the alpha-rhythm, 8–12 Hz) by applying rhythmic visual stimulation at alpha-frequency (10.6 Hz), known to lead to a resonance response in visual areas, and testing its effects on subsequent visual target discrimination. Our data show that rhythmic visual stimulation at 10.6 Hz: 1) has specific behavioral consequences, relative to stimulation at control frequencies (3.9 Hz, 7.1 Hz, 14.2 Hz), and 2) leads to alpha-band oscillations in visual performance measures, that 3) correlate in precise frequency across individuals with resting alpha-rhythms recorded over parieto-occipital areas. The most parsimonious explanation for these three findings is entrainment (phase-locking) of ongoing perceptually relevant alpha-band brain oscillations by rhythmic sensory events. These findings are in line with occipital alpha-oscillations underlying periodicity in visual performance, and suggest that rhythmic stimulation at frequencies of intrinsic brain-rhythms can be used to reveal influences of these rhythms on task performance to study their functional roles.     

The morphochemical characteristics and plastic restructurings of the hippocampal neurons after rhythmic stimulation of an extremity in old rabbits]

The rhythmic activity of CA1 hippocampal neurons was studied after the rhythmic (1-2 Hz) electrocutaneous forelimb stimulation (10-20 min) of 6-84-months-old rabbits. The spectral analysis of neuronal activity revealed the age-dependent decrease in ability to reproduce the rhythm of stimulation. The cytochemical data showed that the protein synthesis in the neurons under study also declined with age.     

Frequency modulation of theta volleys of septal neurons during rhythmic oligosynaptic stimulation in the rabbit

Activity of the neurones with stable theta-bursts was recorded extracellularly in intact and hippocampectomized septum of unanaesthetized chronic rabbits during low-frequency (3-17 Hz) stimulation of horizontal limb of the diagonal band or the lateral septal nucleus. Gradual entrainment and phase-locking of the spontaneous theta-cycles occurred. Two types of entrainment were observed: "entrainment by pause", where interburst interval was reset by the stimuli; and "entrainment by burst", where bursts were time-locked to the stimuli. Such reorganization of the spontaneous bursts occurred in a narrow frequency range of stimulation (from 4 Hz up to 9-12 Hz), with the best resonance following in the range of "basic" theta frequencies of the awake rabbit (5-6 Hz). With stimulation beyond the theta-range three phenomena occurred: shift of the burst frequencies to higher or lower harmonics of stimulation frequencies; complex interactions of basic background frequency with the rhythm of stimulation ("beating"); escape from the influence of the stimuli with return to background theta-burst frequency.     

Modulation of the thalamocortical responses of the sensorimotor cortex during stimulation of the lateral hypothalamus in rabbits in ontogeny

It has been demonstrated that rhythmic stimulation of the relay thalamic nucleus in rabbits at the second half of the first month of postnatal life does not significantly affect the amplitude of the second and third positive components (PC-2 and PC-3) of the positive phase of the thalamocortical responses. Combination of rhythmic stimulation of this nucleus and that of lateral hypothalamic structures is accompanied by a significant increase of the amplitude of both positive components of the thalamocortical responses, the degree of potentiation of PC-3 being higher than that of PC-2. The described potentiation is most evident at the third week of postnatal development of rabbits.     

Postsynaptic neuronal response of a cat sensorimotor cortex during evoked and self-sustained rhythmic "spike and wave" activity

Intracellular correlates of complex sets of rhythmic cortical "spike and wave" potentials evoked in sensorimotor cortex and of self-sustained rhythmic "spike and wave" activity were examined during acute experiments on cats immobilized by myorelaxants. Rhythmic spike-wave activity was produced by stimulating the thalamic relay (ventroposterolateral) nucleus (VPLN) at the rate of 3 Hz; self-sustained afterdischarges were recorded following 8–14 Hz stimulation of the same nucleus. Components of the spike and wave afterdischarge mainly correspond to the paroxysmal depolarizing shifts of the membrane potential of cortical neurons in length. After cessation of self-sustained spike and wave activity, prolonged hyperpolarization accompanied by inhibition of spike discharges and subsequent reinstatement of background activity was observed in cortical neurons. It is postulated that the negative slow wave of induced spike and wave activity as well as slow negative potentials of direct cortical and primary response reflect IPSP in more deep-lying areas of the cell bodies, while the wave of self-sustained rhythmic activity is due to paroxysmal depolarizing shifts in the membrane potential of cortical neurons.I. S. Beritashvili Institute of Physiology, Academy of Sciences of the Georgian SSR, Tbilisi. Translated from Neirofiziologiya, Vol. 18, No. 3, pp. 298–306, May–June, 1986.     

The morphofunctional characteristics of the neurons in the sensorimotor cortex of old rabbits during the trace assimilation of rhythm]

Extracellular neuronal activity was recorded from 460 neurons from alert young (5-7 months), middle-aged (54-65 months) and old (66-85 months) rabbits. Trace rhythmic activity of sensorimotor cortical neurons was examined after long-lasting (10-20 min) rhythmic (0.5-2 Hz) electrocutaneous stimulation of the contralateral forelimb. Spectral analysis of spike activity showed age-related differences in capability of producing a rhythm of previous stimulation in spontaneous neuronal activity. In young animals propriate rhythmic fluctuations of firing rate appeared after the first or second sessions of stimulations (on the first experimental day), in middle-aged ones--after 2-4 sessions (on the second or third days); cortical neurons in old rabbits did not exhibit trace rhythmic activity. Significant morphological changes in glial and neuronal cells were observed in sensorimotor cortex of old rabbits. It is proposed that morphological deteriorations may be the reason of the impairement of trace processes during aging.     

Trace rhythm recruitment by the neurons of the rabbit sensorimotor cortex in old age]

Spectral analysis (ACG and gSP) of the impulse activity of the neurones of the old rabbits sensorimotor cortex allowed to reveal a trace recruitment of the rhythm--CR analogue to time--in after-action f rhythmic stimulation. Connection was established between the number of presented series of periodic electrocutaneous stimulation and expressiveness of the trace rhythm recruitment depending on the animals age. Trace rhythm recruitment took place slower in old animals (54-56 months) than in young ones (up to 1 year), chiefly in 2-3 experimental days after 2-4 series of rhythmic stimulation and was preserved in a small percent of cases the next day after stimulation. In the background activity of a number of neurones an initial periodicity was discovered, which was intensified under the influence of stimulation by another frequency, or the initial rhythm was extinguished, and stimulation rhythm was reproduced. Periodical stimulation in very old animals (66-85 months) practically did not evoke plastic reconstructions of the cortical neurones. Under the influence of the stimulation a non-specific trace increase of the frequency of neurones background activity of the old animals was observed. The revealed characteristics of plastic neurones properties may testify to projected disturbances of mnestic processes at definite age stages of normal aging.     

The structure of the background EEG of the rabbit in the high-frequency portion of the spectrum]

With the purpose of studying the character and structure of high frequency bioelectric activity of rabbits cerebral cortex in the state of calm alertness, the EEG ensembles of different areas of the cortex (sensorimotor, visual, acoustic) and dorsal hippocampus were studied with FFT method. A supposition was made about the presence of systemic organization of the background EEG in rabbits cerebral cortex, reflected, in particular, in the presence of determined components both of chaotic and rhythmic character having different degrees of manifestation. Heterogeneity was revealed in distribution of energies of spectral EEG components in the studied frequency ranges from 14.7 to 100 Hz with predominance of total specific energy value in the band of 14.7-60 Hz. In coherence functions of all the studied pairs of EEG leads rhythmic component, stable in time, was absent. Functions of the mean EEG coherence in the band of 61-100 Hz had significantly greater values in comparison with the values in the band of 14.7-40 Hz.     

Somatosensory evoked potentials in neonates and infants: developmental and normative data

Fifty-two sets of cortical somatosensory evoked potentials (SEPs) were recorded from 23 normal children between the ages of 1 day and 13 weeks with median nerve stimulation. Two bandpass settings 5–1500 Hz and 30–3000 Hz were used; rate of stimulation was 1.1 Hz and sweep-time was 200 msec. The state of wakefulness was documented, but SEPs were obtained and evaluated independently of the child being awake or asleep during the recording. SEPs were present in every recording. The bandpass 30–3000 Hz best differentiated positive and negative early potentials. The bandpass 5–1500 Hz was helpful in some cases, as late slow waves were recorded with this setting. Normative data were established. Mean values were calculated for 3 age groups: 0–2 weeks, 2–6 weeks and 7–13 weeks. P15 and N20 were the first components seen in the newborn, with the P22 becoming the major component by 2–3 weeks of age. The study indicates that maturation of the somatosensory system is fastest during the first 3 weeks of life.     

Functional regulation of the neuronal activity of the rat hippocampus transplanted into the rabbit septum

Embryonal tissue of the rat hippocampus was grafted into dorsolateral septum of the rabbit after its disconnection from the hippocampus. Extracellular investigation of the grafted neurones was performed in chronic conditions 6 to 7 weeks later. The grafted neurones had some characteristics typical of the rat hippocampus (low spontaneous activity and presence of complex spikes, low-frequency rhythmic modulation--below 1 Hz, significant increase of activity level after physostigmine injection). Weak periodic theta-modulation, observed in spontaneous activity of some grafted neurones, became more stable and appeared in additional units after injection of physostigmine, under the influence of electrical stimulation of the septum and reticular formation and in response to sensory stimulation of the host animal. Its frequency was 4.5-6.0 Hz, as in the host septum. Microelectrode investigation for 5-10 days provoked lymphocyte infiltration of the grafts, which was not observed in the grafts not subjected to such treatment.     

The effect of a vasopressin analog on the reaction of the command neurons in the defensive behavior of the edible snail during nerve stimulation

On the level of snail command neurones of the defensive behaviour an increase was revealed of the amplitude of summary EPSP recorded in command neurones in response to nerve stimulation, an increase of probability of appearance of action potential in the reaction and, under certain conditions, a significant decrease of habituation speed at rhythmic (0.1 Hz) nerve stimulation against the background of peptide. The latter effect was found during comparison of groups of neurones--control neurones and those to which the peptide was presented before the first series of stimulation in the test, i. e. without preliminary elaboration of habituation. The decrease of the speed of habituation concerned both the amplitude of summary EPSP and the probability of action potential appearance in the reaction. All these changes against the background of vasopressin analogue may be the basis of the increase of spike reactions of command neurones of the defensive behaviour and thus the basis of the initiation or the increase of behavioural defensive reactions. The obtained effects were not protracted and took place only in the presence of the peptide in the extracellular medium.     

Regulation of indoleamine 2,3-dioxygenase activity in the small intestine and the epididymis of mice

Indoleamine 2,3-dioxygenase activity was found to be ubiquitously distributed in various tissues of mice, such as brain, lung, stomach, intestine, and epididymis. The highest enzyme activity was detected in the alimentary canal and the epididymis. Developmental and daily rhythmic changes of indoleamine 2,3-dioxygenase activity and the effects of various regulatory factors were studied with the supernatant fractions derived from the small intestine and the epididymis. The enzyme activity in these two tissues was absent during the first 2 weeks (the weaning period). From the third week, there was a rapid increase in activities and a maximum was reached when the mice were 8 to 10 weeks of age (adolescence). The enzyme activity in the small intestine then gradually diminished to zero level at 30 weeks of age (prime) or later, while that in the epididymis remained at the high level throughout 69 weeks of age (senescence). The enzyme activity of the small intestine from mice fed during the hours 9:00–13:00 showed daily rhythmic changes; high in the daytime and low at night. Under night feeding (21:00–1:00), the enzyme activity was high at night and low in the daytime. The epididymal enzyme activity showed no daily fluctuations by either feeding schedule. With regard to the developmental and daily rhythmic changes, indoleamine 2,3-dioxygenase activity in the small intestine was similar to that of hepatic tryptophan 2,3-dioxygenase. However, in contrast to the hepatic tryptophan 2,3-dioxygenase activity, indoleamine 2,3-dioxygenase activity in the small intestine and the epididymis was not affected by adrenalectomy or intraperitoneal administration of adrenal steroid or tryptophan.     

Dynamic characteristics of the human resonance EEG responses to rhythmic photostimulation

The development of the resonance EEG responses of the left and right occipital areas was studied in right-handed men during prolonged (12 or 120 s) rhythmic, photostimulation with the intensity of 0.7 J and frequencies of 6, 10, and 16 Hz. Analysis of the EEG fine spectral structure was applied to compare the accumulated baseline EEG spectra and EEG spectra during photostimulation, to observe the dynamics of the short-term spectra and to detect power changes in the EEG narrow spectral band sharply coincident with the stimulation frequency. The more pronounced EEG responses to photostimulation were observed in subjects with the initially low EEG baseline, α-rhythm. Two-minute flash trains produced a substantial increase in the EEG power within the stimulation frequency with superposed oscillatory processes with different periods. These fluctuations are considered a reflection of intricate interaction between the adaptive and resonance EEG responses to the presented intermittent stimulation. Under 12-s stimulation the resonance EEG responses are steadily recorded within the first 3 s of stimulation and immediately after the flash cessation EEG power at the stimulation frequency returns to the initial level. The resonance EEG responses were more pronounced in the right hemisphere than in the left one, especially, at the stimulation frequencies of 6 and 16 Hz. With increasing the stimulation frequency, the maximum of resonance EEG responses was reached earlier. Under the stimulation frequency of 6 Hz, the maximal response was recorded 9–12 s after the beginning of flashes, at the frequencies of 10 and 16 Hz, it was recorded within 3–6 and 3 s, respectively.     

Theta driving both inhibits and potentiates the effects of nicotine on dentate gyrus responses

The medial septal area of conscious rats was stimulated through previously implanted electrodes at a frequency of 7.7 Hz for 20 min each day for 7 days to evoke rhythmic slow activity in CA1 at a similar frequency to spontaneous theta. Two weeks later in the anaesthetized rats the effects of a single subcutaneous injection of nicotine (0.4 mg x kg(-1)) on fEPSPs, evoked in the dentate gyrus by separate stimulation of the MPP and LPP, were studied and compared with those obtained in controls. Nicotine increased the firing of locus coeruleus neurones and the slope of the fEPSPs evoked by LPP stimulation, but not by MPP stimulation. Prior theta driving considerably increased the effect of nicotine on the responses evoked by stimulation of the MPP and abolished the nicotine-induced potentiation of the responses evoked by stimulation of the LPP. The results are attributed to theta driving increasing the amount of noradrenaline released by nicotine and to noradrenaline producing a beta-adrenoceptor long-lasting potentiation at the medial perforant path synapse and a long-lasting depression at the lateral perforant path synapse.     

Investigation of the functional role of the septum

A spectral-correlation analysis was made of the EEG recorded from the motor and visual cortex, hippocampus, and septum (in some series of experiments, the lateral geniculate body and mesencephalic reticular formation also) of rabbits in the presence or absence (background) of sensory stimulation. To investigate the functional role of the septum as a pacemaker the method of "rhythm binding" by electrical stimulation of the septum (lateral and medial nuclei) was used. By electrical stimulation of the medial nucleus of the septum at a frequency of 4–30 Hz rhythm binding was successfully obtained in all regions studied. Maximal rhythm binding was observed in cortical potentials. It is postulated that cortical rhythmic activity is generated as a result of physiological interaction between cortex and septum.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 8, No. 3, pp. 267–275, May–June, 1976.     

Ontogenetic analysis of the EEG-activation reaction in children

Frequency characteristics of EEG-reaction have been analysed in children of 3-10 years in the process of realization of the orienting reflex to sound stimuli. An enhancement of theta- and alpha-oscillations has been found in such conditions which becomes less pronounced to 9-10 years. In the last age group, the reaction of blockade of the basic rhythm of the background EEG (alpha-oscillations of 10 Hz) is formed, combined with local enhancement of other rhythmic components of this frequency range. Functional significance of different forms of EEG activation reaction has been determined. The question is discussed of functional heterogeneity of rhythmic components of the alpha-range and their differential involvement in the reaction reflecting complex mechanisms of cortical activation.     

The characteristics of trace rhythm reproduction by the neurons of the rabbit sensorimotor cortex in the aftereffect of periodic stimulation]

Trace rhythm recruitment (TRR)--CR analogue to time was studied appearing in response to prolonged electrocutaneous stimulation of the forelimb of the alert rabbit with the frequency 0.5-1-2 Hz. The activity was recorded of 180 cells of the sensorimotor cortex before (80) and after (100) periodical stimulation during 10-20 min. The first series of rhythmic stimulation led to a short-term TRR of the stimulation frequency, the following series formed a clear TRR, preserved for several days. The possibility was revealed of "relearning" of neurones at stimulation rhythm change. The ability of TRR phenomenon of extinguishment, prolonged preservation and reproduction of traces, "relearning" brings it nearer to the processes, analogous to the temporal connection. The ability to reveal distinctly and to quantitatively estimate the characteristics of the applied stimulus fixated by the neurones, makes this model perspective for comparable study of the memory traces at the neuronal level in the animals of various ages.     

Self-sustained rhythmic spike and wave activity and response of glial and nerve cells in the cat sensorimotor cortex

During acute experiments on unanesthetized cats, immobilized with myorelaxants, it was found that during rhythmic stimulation (8–14 Hz, duration: 10 sec) of the ventroposterolateral thalamic nucleus brief hyperpolarization is succeeded by depolarization in the pyramidal neurons of the sensorimotor cortex. Following this depolarization, rhythmic (approximately 3 Hz) paroxysmal depolarizing shifts in membrane potential are produced by ending stimulation, succeeded by protracted hyperpolarization and termination of rhythmic wave activity. Depolarization only is observed in glial cells, however, while hyperpolarization sets in after hyperpolarization is completed in the neurons. It is suggested that long-term changes in the membrane potential of cortical cells could make some contribution to the setting up and termination of rhythmic spike and wave activity.I. S. Beritashvili Institute of Physiology, Academy of Sciences of the Georgian SSR, Tbilisi. Translated from Neirofiziologiya, Vol. 18, No. 3, pp. 319–325, May–June, 1986.