Relationship between posterior thalamic nucleus unit activity and parietal cortical rhythms (beta) in the waking cat.

Cat behavioural states of attentive fixation on a target are associated with episodes of electrocortical rhythms at 40 Hz ("beta rhythms") in the parietal cortex. Previous field potential studies indicate that the nucleus posterior pars medialis of the thalamus displays this particular rhythmic activity. We investigated single units of the nucleus posterior pars medialis and its surrounding nuclei to assess their participation in the cortical beta rhythms. Only a small proportion of thalamic cells underwent changes in their firing pattern during beta episodes. "Beta-related cells" were localized in the nucleus posterior pars medialis or its immediate vicinity; no such beta-related cells were found in other regions of the lateral thalamus. Some beta-related cells showed a one-spike to one-wave relationship ("homorhythmicity"), while others displayed a prolonged decrease or a suppression of their firing throughout each beta episode ("pause cells"). For comparison, neurons in the same thalamic area were also recorded during sleep episodes with slow waves and spindles: there was no correlation between spindles and cell firing. Thus, the nucleus posterior pars medialis thalamic nucleus contains cells whose firing is correlated with the beta rhythms. No such correlation was found with sleep spindles.

Anatomical localization of cortical beta rhythms in cat.

Beta electrocorticographic rhythms (40 Hz) develop during motionless focused attention in two distinct cortical foci in cats. A cytoarchitectonic study was performed to determine the precise location of these foci. Electrode tips recording beta rhythms were found: (i) in motor areas 4 gamma and 6a beta, in a band extending from the postcruciate cortex to the walls of the presylvian sulcus, crossing the frontal pole (anterior beta focus); (ii) in the posterior parietal associative area 5a, along the divisions of the ansate sulcus, extending to the mesial aspect of the hemispheres (posterior beta focus). The two foci are separated by areas 3, 2 and 1, where beta rhythms were never recorded. The fact that both these areas, containing giant pyramidal cells, develop a specific type of activity during immobility may have a functional meaning: area 5 may be involved in the cat as it is in the monkey in the control of motor behaviour.

Effects of two neuroleptic drugs on focal somatoparietal rhythms in free awake cats.

In freely moving cats, systemic administration of Chlorpromazine (CPZ) and Haloperidol (HP), two neuroleptics which presumably block catecholamine receptors, leads to a suppression of the waking mu-type rhythms. There is a clear difference, however, in that CPZ leads to slow wave sleep while HP only induces sustained drowsiness. The difference in their action is tentatively explained by their differential effect on an enzyme that is not involved in the metabolism of biogenic amines, tryptophan pyrrolase. We suggest that it is through this pathway that CPZ might increase and HP might decrease brain tryptophan and thereby lead to opposite changes in brain 5-HT content.

Modification of spontaneous ECoG and behavior in cat by monoamine precursors.

The effects of increasing the brain monoamine content on the three types of localized rhythms identified in somatic area I ("mu-type rhythms") were studied in freely moving cats. Increases in brain monoamines (catecholamines and serotonin) were produced by injecting L-Dopa and 5-HTP, L-tyrosine, L-tryptophan and tryptamine were also tested. It is concluded that the two subsets of rhythms, one accompanying high levels of alertness and vigilance and the other appearing during quiet wakefulness, are related to the catecholaminergic systems, while those characterizing drowsiness seem to be controlled by a serotoninergic system. The latter, however, is distinct from that controlling slow sleep.

Selective modification of spontaneous ECoG rhythms of the cat somesthetic cortex by psychoactive drugs: behavioral correlates.

Three psychoactive drugs with known central effects were administered to the freely moving cat in order to study their action on spontaneous rhythmic activities recorded from the primary somesthetic cortex, which are analogous to the rolandic mu rhyhm in man. The ECoG patterns obtained are qualitatively identical to those of the normal subject, but their temporal organization is profoundly disturbed by the action of the drugs. The normal ECoG consists of three rhythmic systems with distinct frequencies and displays a considerable time variability. In contrast, psychoactive drugs induce a stabilized pattern with only one type (or at most two types) of rhythm prevailing for one or several hours, which never occurs under normal conditions. These ECoG rhythms underlie various behavioral states. Under d-amphetamine, correspondence remains excellent between behavior and ECoG; under Ditran, complete dissociation occurs; finally, LSD represents a borderline case in which ECoG and behavior are partially correlated and partially dissociated.

Action of tianeptine on focalization of attention in cat.

Tianeptine, an antidepressant substance devoid of sedative action, was investigated for its effects on focalization of attention in cats, using combined behavioural and electrocorticographic (ECoG) observations. The ECoG index was the presence of 40 Hz frontoparietal rhythmic cortical activities, developing while the animal displayed a behaviour suggesting focused attention. Cats were observed in two conditions: a "neutral" one, with no specific target, and another one, "of focused attention" where a live mouse placed in a transparent box was present in the recording room, each test lasting for 90 min. After treatment the animals displayed increased attention even in the neutral situation, where the cat, instead of sleeping like during control sessions, payed sustained attention to its environment. Conversely, at corresponding doses, amitriptyline, another antidepressant drug chosen as a reference, induced uninterrupted slow sleep even in the situation of focused attention.

Effect of DSP4, a neurotoxic agent, on attentive behaviour and related electrocortical activity in cat.

Six behaving cats were administered N-(2-chloroethyl)-N-ethyl-2-bromobenzylamine (DSP4) (i.p. 10 mg/kg), a neurotoxic agent known to destroy noradrenergic endings in the CNS. They were placed, both before (control) and after treatment in 3 different situations, each time for 90 min: (1) a 'neutral' one (N) with no significant stimuli; (2) another one eliciting focused attention (F); (3) a third one, creating a situation of 'expectancy of an event to occur' (E), with the animal usually displaying a posture of quiet waking. Simultaneously, the electrocorticogram (ECoG) was recorded from the sensorimotor and parietal cortex through implanted electrodes. We already knew and were able to confirm that the normal animals (i.e. before treatment) displayed distinct behavioural patterns depending on the situation and distinct accompanying parietofrontal ECoG activities, with a dominance of drowsiness and sleep during N, that of a 36-Hz 'beta' rhythms in condition F, and of 14-Hz 'mu' rhythms in condition E. It was shown that the prevailing attitude of the animals after treatment was now, in all 3 situations, that of 'quiet waking and/or expectancy-like watching', with a large if not exclusive dominance of only one ECoG pattern, namely mu. These changes were considered as due to a release of the mu system from a noradrenergic modulatory blockade, in accordance with some of our previous data. An immunohistochemical study with anti-tyrosine hydroxylase antibody was also performed; it confirmed that after DSP4 treatment there were substantial alterations in the immunoreactivity of locus coeruleus cells, the structure which is likely to be involved in this NAergic control of the mu rhythms and of its concomitant behaviour.

Ventral mesencephalic tegmentum (VMT) controls electrocortical beta rhythms and associated attentive behaviour in the cat.

When a cat is immobile, very alert and displaying behaviour suggesting focused attention toward a target in its environment, beta rhythms (ca. 40 Hz) develop in the fronto-parietal cortical areas. After bilateral electrolytic lesions of the ventral mesencephalic tegmentum (VMT), these beta rhythms are suppressed (while other cortical activities, with other behavioural correlates, persist), and at the same time, attentive immobility is no longer observed: the same experimental situation as in the control now elicits locomotor hyperactivity. Arguments are produced, favouring the hypothesis that both behavioural immobility and the accompanying thalamocortical beta rhythms are controlled through one of the dopaminergic system that originate from the VMT and are distinct from the nigrostriatal one.

Effects of locus coeruleus lesions on vigilance and attentive behaviour in cat.

Previous data have suggested that in the cat, expectancy behaviour (waiting for a target to appear) and associated electrocortical, focal, synchronized activity ('mu' rhythms) are modulated by a noradrenergic system possibly originating from the locus coeruleus (LC). To test the latter hypothesis, we have examined the behavioural and ECoG changes induced after bilateral LC lesions. Our results demonstrated that destruction of the anterior 3/4th of the LC (A6 noradrenergic cell group) resulted in a considerable increase of mu rhythms and expectancy behaviour, without episodes of drowsiness that normally occur. Destruction of the posterior fourth of LC (A4 noradrenergic group) only increased the duration of slow sleep. Extending the A6 lesion to include the dorsal ascending noradrenergic bundle also increased the expectancy behaviour and mu rhythms. Finally, when the nucleus subcoeruleus was also involved, the duration of slow sleep and the frequency of paradoxical sleep episodes increased. These findings indicate that the LC exerts an inhibitory effect on structures involved in the induction and persistence of expectancy behaviour with accompanying mu rhythms.

Effects of mediodorsalis thalamic nucleus lesions on vigilance and attentive behaviour in cats.

In this study performed on 10 cats we analysed the effects of limited lesions of nucleus medialis dorsalis of the thalamus (MD) on behaviour and on some specific electrocorticographic (ECoG) patterns, known from previous works to accompany various states of waking attentiveness. The animals were tested during 90 min in three distinct behavioural situations; a neutral one (NS) where they simply explored their environment and then usually went to sleep; a second one (FA) where they could watch a mouse (but not catch it), which favoured 'focussed' attentiveness accompanied by the development of rhythmic ECoG activities in the anterior frontoparietal cortex ('beta rhythms' at 40 Hz); a third one (EX) with a hidden mouse whose appearance the cat was waiting for ('expectancy') which rather than favouring the development of beta rhythms elicited the appearance of another rhythmic activity dominating in somatic area SI, 'mu rhythms' at 14 Hz. The duration of each waking behaviour and its ECoG concomitant as well as that of slow wave and of paradoxical sleep were compared in each cat before and after lesion. Our results showed that MD lesions situated in the posterior part of the nucleus tended to increase the time occupied by focussed attentive behaviour and the accompanying beta rhythms. On the other hand, anteriorly located MD lesions elicited a concomitant reduction of both manifestations, behavioural and electrocortical. No such contrasting effects could be systematically noticed for the durations of expectancy, of slow sleep and of paradoxical sleep. These findings were discussed considering previous data showing that at least two systems project upon MD, that play distinct--in a way even antagonistic--roles in focussed attention upon a target. Both originate from the ventral tegmental mesencephalic area; one reaches MD through the ventral striatum (nucleus accumbens), the other one through the amygdala. The MD nucleus thus receiving contrasting information may participate in a final adjustment of the attentive state of the animal to its environment.

Noradrenaline-like terminals in the cat nucleus ventralis posterior of the thalamus.

Noradrenaline-like immunoreactivity in the cat nucleus ventralis posterior of the thalamus was investigated using an indirect immunocytochemical technique. Specific antinoradrenaline antibodies, raised in rabbits, were used. It was first verified that these antibodies recognize noradrenaline cells bodies of the locus coeruleus and their ascending axons in the ascending noradrenergic tract. In the nucleus ventralis posterior itself, noradrenaline-like fibers were observed. They were either randomly distributed or grouped around nonlabeled cell bodies. These neurons were generally oblong and measured 60-80 microns. With electron microscopy, preliminary results showed immunoreactive fibers in close apposition to unlabeled cell bodies or dendrites. The precise nature of these profiles was sometimes difficult to ascertain, since experiments were done in presence of detergent. In some cases symmetric synapses might be observed between immunoreactive axon terminals and unlabeled dendrites. The specificity of the reaction is discussed in the light of several control experiments.

Somatosensory rhythms in the awake cat: a single unit exploration of their thalamic concomitant in nucleus ventralis posterior and vicinity.

A microelectrode study was performed in n. ventralis posterior (VP) of the fully alert cat, to study the correlation between thalamic unit activity and the cortical synchronized 14 c/sec rhythms that develop in the somatic area I for forepaw and wrist projections, when the animal is in a state of "quiet waking". (i) Only a small proportion of VP cells underwent changes during the studied cortical rhythms. (ii) None of these cells were typical thalamo-cortical relay cells carrying tactile messages to the cortex. (iii) Cells altering their discharge were of two types, rhythmic (R) cells, discharging at the frequency of the cortical rhythms, and tonic (T) cells, displaying an overall, sustained change during the whole sequence of cortical 14 c/sec. (iv) Among R cells, some were long axon thalamo-cortical cells and others, likely to be interneurones. (v) Some T cells increased their firing rate during rhythmic trains, others came into silence during the same period. (vi) The thalamic circuitry responsible for this particular set of "quiet waking rhythms" is discussed.

Nucleus reticularis thalami participates in sleep spindles, not in beta rhythms concomitant with attention in cat.

In the awake cat, characteristic electrocortical synchronized oscillations (beta rhythms) can develop on the parietal cortex in specific behavioural situations such as during attentive fixation of a target. These activities differ from other synchronized rhythms which accompany slow sleep ("spindles") and are more widespread. It is shown here through single unit recording from the thalamic nucleus reticularis (RET), that the latter structure participates in spindles (as already postulated by other authors), but not in the beta rhythms which seem to depend on a more restricted thalamic focus in the posterior thalamic nucleus. These data thus support the idea that RET plays a role in slow-wave sleep but not in the cognitive operations involved in focussing attention.

Comparative anxiolytic and hypnotic effects of 3 benzodiazepines on baboons.

The ECoG and behavioural effects of administering 3 benzodiazepines (diazepam, nitrazepam, lorazepam) were tested in baboons restrained in a chair, with arms fastened (a situation which is considered 'anxiogenic'). Considering our previous data with ECoG recording of focal rhythms from area somatic I, the 3 drugs were revealed as 'anxiolytic'. Differences were noticed with respect to their hypnotic effects: no sleep was elicited under diazepam; a considerable amount of slow sleep was observed with lorazepam; nitrazepam produced a succession of short waking/sleep cycles.

Alpha rhythm in the cat thalamus.

In cats, electrocortical rhythms at about 10 Hz displaying common characteristics with the human alpha rhythm [1] were recorded from the part of the visual cortex that includes the anterior half of areas 17 (on the cortical convexity), of area 18 and of their common limit, representing the projection of the lower contralateral visual quadrant and that of the lower vertical meridian. It is shown here that activities highly correlated with these rhythms and at the same frequency, were recorded from the anterior half of the laminar dorsal lateral geniculate body (mediodorsal part of layer A), and also from a limited area medial to this nucleus. The cat thalamocortical alpha system thus appears to concern only the projection of the lower quadrants, probably excluding the area centralis itself.

A 10 Hz "alpha-like" rhythm in the visual cortex of the waking cat.

Rhythms at about 10 Hz were recorded from the primary visual cortex of the cat (anterior part of area 18), with characteristics close to those of the alpha rhythm in man: frequency band (7-14 Hz), localization and reactivity to visual stimulation. Coherence analysis of this activity with the "mu" rhythms on the somatosensory cortex showed that although both types develop in the same overall behavioural situations (quiet waking and/or expectancy of an event to occur), they are independent.

[Unit activity of neurons of the posterior ventral nucleus of the thalamus for various waking stages in the normal cat]. / Activité unitaire de neurones du noyau ventral postérieur du thalamus pour divers degrés de vigilance chez le chat normal.

Using the electrocortical activity in the sensorimotor cortex as an index, three distinct levels of motionless waking can be identified in the cat, all three different from active waking and from slow sleep (attentive waking, quiet waking and drowsiness). It has now been shown that spontaneous and evoked single unit activities in n. ventralis posterior of the thalamus undergo significant changes when passing from one level of waking into another one.