Unit activity of the reticular and relay nuclei of the cat thalamus

Relations between neurons of the reticular and specific relay nuclei of the thalamus were studied in cats immobilized with tubocurarine. Under the influence of stimulation of the reticular nucleus (RN) unit activity in the thalamic relay nuclei was found to be considerably modulated. Cases of the appearance of IPSPs (possibly of monosynaptic nature), evoked by stimulation of RN, in neurons of the ventroposterolateral nucleus (VPLN) and lateral geniculate body (LGB) are described. During simultaneous recording of unit activity in RN and VPLN or LGB by means of two electrodes interaction of several types was found: inhibition of discharges of VPLN or LGB neurons accompanied by excitation of RN neurons: alternation of excitation-inhibition in neuron pairs in RN and VPLN or RN and LGB during low-frequency afferent or cortical stimulation (in this case excitation of RN neurons is associated with inhibition of VPLN or LGB neurons), and strengthening of the discharge of VPLN or LGB neurons during excitation of RN neurons. The possibility of the existence both of direct monosynaptic inhibition of activity of VPLN or LGB relay neurons under the influence of excitation of RN neurons and of their inhibition by activation of hypothetical interneurons of the relay nuclei themselves is accepted.I. S. Beritashvili Institute of Physiology, Academy of Sciences of the Georgian SSR, Tbilisi. Translated from Neirofiziologiya, Vol. 13, No. 1, pp. 24–31, January–February, 1981.     

Effect of deafferentation of the lateral geniculate body on its background activity

Elimination of reticular inputs to the lateral geniculate body (LGB) by sectioning of one half of the midbrain operculum, did not affect significantly the characteristics of the LGB evoked potential to light stimulus. At the same time LGB response to stimulation of the reticular formation by a single current impulse, though did not disappear completely, but changed greatly: its latency became twice as long, the negative component of the response was no more recorded. In conditions of LGB deafferentation, the characteristics of all rhythms of its electrical activity, besides the alpha-like one, considerably changed. At the same time, exactly this last rhythm underwent the greatest changes on the EEG of the visual cortex. On the basis of the obtained data it is suggested that the reticular formation takes a considerable and multiple part in generation of LGB rhythmic activity and that changes in its characteristics are clearly reflected in the ECoG rhythms formation. Retention of the LGB visual evoked potential and of the response to stimulation of the reticular formation after the section of one half of the midbrain operculum testifies to the presence of several reticular inputs to LGB.     

Ascending and descending efferent pathways of the midbrain reticular formation of the cat (radioautographic study)

By means of the anterograde axoplasmic transport technique for a mixture of labelled aminoacids (3H-leucine and 3H-proline), ascending and descending systems of the reticular formation fibers in the cat mesencephalon have been studied. Projections from the mesencephalon reticular formation (MRF) ascend to the subthalamus, lateral, dorsal and periventricular hypothalamus, to the periventricular nuclei of the midline and to the intralaminar nuclei of the thalamus. The descending pathways project to the grey substance surrounding the aqueduct of cerebrum, locus coeruleus, parabrachial region and reticular formation of the pons and medulla oblongata. The projections to the reticular nucleus of the thalamus, ventral nucleus of the external geniculate body and superior colliculi arise from the dorsal half of the MRF, and projections to the striatum, lateral reticular nucleus of the medulla oblongata--from its ventral half. Most of the structures are reciprocally connected with the MRF.     

Interaction of reticular structures in the brain of the cat

In acute and chronic experiments on 35 cats an inhibitory influence was found of the caudal reticular nucleus of pons Varolii on unit activity of the sensorimotor cortex and dorsal part of the midbrain reticular formation. The influence of this structure on unit activity of the ventral part of the midbrain reticular formation was mainly of a facilitatory character. Activation of the ventral part inhibited the unit activity of the dorsal part of the same structure. Consequently, the caudal reticular nucleus of pons Varolii elicits inhibition at the level not only of the cerebral cortex but also of the midbrain reticular formation (of its dorsal part). The character of these influences coincides with that of unit activity changes of these two areas of the midbrain reticular formation during the development of the paradoxical phase of sleep. The obtained facts must underlie the stopping of convulsive activity in this phase of sleep.     

Chronic pain as a reticular formation syndrome

Evidence was previously presented to support the thesis that chronic pain is activated by neuronal elements that make up the multisynaptic short axon core of the reticular system (Andy and Peeler 1985). The present thesis, that chronic pain is a reticular formation syndrome, is based on a retrospective analysis of four patients with chronic pain who were successfully treated with a lesion in the anterior thalamus and stimulation electrode implants in the posterior thalamus and pontomesencephalic brain stem. The reticular formation was the common underlying anatomic substrate at those three sites. In addition to chronic pain, all the patients had other symptoms attributable to other body organs and systems. The number and type of symptoms that made up the syndrome differed between patients. Symptoms making up the core of the syndrome were pain, anxiety, nervousness, insomnia, and depression. Experimental and clinical findings are briefly presented to demonstrate the various reticular formation sites, pragmatically considered "reticular functional systems," from which symptoms may arise. It is hypothesized that the symptoms are recruited by a low threshold "pain oscillator" that is generated at one reticular site and subsequently permeates the rest of the reticular system. Therapeutic stimulation inactivates the low threshold system by "jamming" it.     

Common system of sparsely branched projection (reticular) NADPH-diaphorase neurons in the human forebrain formations built from densely branched cells

Cell morphometry with statistical analysis (using 9 parameters) of densely branched projection and sparsely branched reticular neurons was performed in the human forebrain formations built from densely branched projection neurons (the entorhinal cortex, striatum, nucleus accumbens basolateral amygdala, and dorsal thalamus). The reticular neurons included scattered reticular neurons and marginal reticular neurons of the dorsal thalamus. Golgi method and staining for NADPH-diaphorase were used. The scattered reticular neurons of different formations under study did not differ in any of the 9 parameters, whereas they significantly differed from the main projection neurons in 5 to 7 parameters (except one comparison with the difference in 2 parameters). Within the same formation, the scattered reticular and main projection densely branched neurons differed in 7 to 9 parameters. The endbrain scattered reticular neurons expressed NADPH-diaphorase, while in the dorsal thalamus only the medium marginal reticular neurons were NADPH-diaphorase-positive. Thus, a common system of ancient integrative reticular neurons expressing NADPH-diaphorase exists in the examined human forebrain formations. The evidence obtained by us and the literature data point to the projection nature of the scattered reticular neurons (to the V and VI neocortical layers), which suggests their modulatory influence on descending neocortical pathways.     

A thalamic reticular networking model of consciousness

[Background]  

It is reasonable to consider the thalamus a primary candidate for the location of consciousness, given that the thalamus has been referred to as the gateway of nearly all sensory inputs to the corresponding cortical areas. Interestingly, in an early stage of brain development, communicative innervations between the dorsal thalamus and telencephalon must pass through the ventral thalamus, the major derivative of which is the thalamic reticular nucleus (TRN). The TRN occupies a striking control position in the brain, sending inhibitory axons back to the thalamus, roughly to the same region where they receive afferents.     

Postnatal development of thalamic reticular nucleus projections to the anterior thalamic nuclei in rats

The thalamic reticular nucleus (TRN) projects inhibitory signals to the thalamus, thereby controlling thalamocortical connections. Few studies have examined the development of TRN projections to the anterior thalamic nuclei with regard to axon course and the axon terminal distributions. In the present study, we used parvalbumin (PV) immunostaining to investigate inhibitory projections from the TRN to the thalamus in postnatal (P) 2- to 5-week-old rats (P14–35). The distribution of PV-positive (+) nerve fibers and nerve terminals markedly differed among the anterior thalamic nuclei at P14. Small, beaded nerve terminals were more distributed throughout the anterodorsal nucleus (AD) than in the anteroventral nucleus (AV) and anteromedial nucleus (AM). PV+ fibers traveling from the TRN to the AD were observed in the AV and AM. Nodular nerve terminals, spindle or en passant terminals, were identified on the axons passing through the AV and AM. At P21, axon bundles traveling without nodular terminals were observed, and nerve terminals were distributed throughout the AV and AM similar to the AD. At P28 and P35, the nerve terminals were evenly distributed throughout each nucleus. In addition, DiI tracer injections into the retrosplenial cortex revealed retrogradely-labeled projection neurons in the 3 nuclei at P14. At P14, the AD received abundant projections from the TRN and then projected to the retrosplenial cortex. The AV and AM seem to receive projections with distinct nodular nerve terminals from the TRN and project to the retrosplenial cortex. The projections from TRN to the AV and AM with nodular nerve terminals at P14 are probably developmental-period specific. In comparison, the TRN projections to the AD at P14 might be related to the development of spatial navigation as part of the head orientation system.Key words: Thalamic reticular nucleus, parvalbumin, axon terminal, development, anterior thalamic nucleus, rat     

Evolution of the reticular formation

The reticular formation of mammals contains numerous nuclei which can be recognized by their projection patterns, cytoarchitectonics, and neuropeptide/neurotransmitter content. We have identified reticular nuclei in representatives from numerous reptilian groups and ascertained presence or absence of these reticular nuclei in an attempt to use neuronal occurrence as a tool to determine phylogenetic relationships. Recently these studies have been extended to two elasmobranchs, a galeomorph shark and a ray. In this report, we concentrate on three medullary spinal projecting reticular nuclei, reticularis gigantocellularis, reticularis magnocellularis, and reticularis paragigantocellularis. We found that all three nuclei were present in rats, lizards, and elasmobranchs, but one nucleus was absent in crocodilians, and two nuclei were absent in turtles. Thus brain organization may give us clues to phylogenetic relationships. Moreover, these three reticular nuclei exhibited remarkably similar cellular morphology in mammals, reptiles, and elasmobranchs.     

Single unit responses of the reticular and ventral anterior nuclei of the thalamus to electrical stimulation of the centrum medianum

In acute experiments on cats anesthetized with thiopental (30–40 mg/kg, intraperitoneally) and immobilized with D-tubocurarine (1 mg/kg) responses of 145 neurons of the reticular and 158 neurons of the ventral anterior nuclei of the thalamus to electrical stimulation of the centrum medianum were investigated. An antidromic action potential appeared after a latent period of 0.3–2.0 msec in 4.1% of cells of the reticular nucleus and 4.4% of neurons of the ventral anterior nucleus tested in response to stimulation. The conduction velocity of antidromic excitation along axons of these neurons was 1.7–7.6 m/sec. Neurons responding with an antidromic action potential to stimulation both of the centrum medianum and of other formations were discovered, electrophysiological evidence of the ramification of such an axon. Altogether 53.8% of neurons of the reticular nucleus and 46.9% of neurons of the ventral anterior nucleus responded to stimulation of the centrum medianum by orthodromic excitation. Among neurons excited orthodromically two groups of cells were distinguished: The first group generated a discharge consisting of 6–12 action potentials with a frequency of 130–640 Hz (the duration of discharge did not exceed 60 msec), whereas the second responded with a single action potential. Inhibitory responses were observed in only 0.7% of neurons of the reticular nucleus and 4.4% of the ventral anterior nucleus tested. Afferent influences from the relay nuclei of the thalamus, lateral posterior nucleus, and motor cortex were shown to converge on neurons responding to stimulation of the centrum medianum.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 1, pp. 36–45, January–February, 1980.     

Differentiation of efferent projections of the medial (cuneiform nucleus) and lateral regions of the reticular formation in cat midbrain

Efferent connections of medial (nucleus cuneiformis) and lateral regions of the midbrain reticular formation (MRF) were investigated using an anterograde autoradiographic technique in cats. Efferent fibers from the MRF ascend to the globus pallidus, substantia innominata, hypothalamus, subthalamus, and nonspecific associative and relay nuclei of the thalamus. Descending pathways to the conclusion that the cuneiform nucleus is more of a nonspecific structure than an association auditory center. The lateral reticular region had numerous projections to the lateral geniculate body and, together with the parabigeminal nucleus, forms the midbrain visual complex.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 17, No. 5, pp. 646–652, September–October, 1985.     

Early lesion of the reticular thalamic nucleus disrupts the structure and function of the mediodorsal thalamus and prefrontal cortex

The thalamic reticular nucleus (TRN), part of the thalamus, is a thin GABAergic cell layer adjacent to the relay nuclei of the dorsal thalamus. It receives input from the cortex and other thalamic nuclei and provides major inhibitory input to each thalamic nucleus, particularly the mediodorsal nucleus (MD). As the MD is important for supporting optimal cortico–thalamo–cortical interactions during brain maturation, we hypothesized that that early damage to the TRN will cause major disturbances to the development and the functioning of the prefrontal cortex (PFC) and the MD. Rat pups at P4 were randomized in three groups: electrolytic lesion of TRN, TRN‐sham‐lesion group, and the classical control group. Seven weeks later, all rats were tested with several behavioral and cognitive paradigms, and then perfused for histological and immunohistochemical studies. Results showed that TRN lesion rats exhibited reduced spontaneous activity, high level of anxiety, learning and recognition memory impairments. Besides the behavioral effects observed after early TRN lesions, our study showed significant cytoarchitectural and functional changes in the cingulate cortex, the dorsolateral and prelimbic subdivisions of the PFC, as well as in the MD. The assessment of the basal levels of neuronal activity revealed a significant reduction of the basal expression of C‐Fos levels in the PFC. These experiments, which are the first to highlight the effects of early TRN lesions, provided evidence that early damage of the anterior part of the TRN leads to alterations that may control the development of the thalamocortical–corticothalamic pathways.     

Afferent connections of the basolateral division of the amygdaloid complex of the cat brain

Injection of horseradish peroxidase into the basal macrocellular and lateral nuclei of the amygdaloid complex (BLAC) in the cat brain has revealed their rich thalamic afferentation. On the BLAC there are massive projections of: a) nuclei of the middle line of the precommissural pole of the dorsal thalamus (anterior parts of the paratenial, interanteromedial and reunial nuclei), as well as the whole anterior paraventricular nucleus, medial part of the ventral posteromedial nucleus; b) postcommissural nuclei of the dorsal thalamus; some "nonacustical" nuclei of the internal geniculate body (ventrolateral nucleus, medial and macrocellular parts and the most caudal end of the internal geniculate body). Rather essential are projections of the "posterior group nuclei", those of the suprageniculate nucleus, of some parts of the ventral thalamus (subparafascicular nucleus, marginal and peripeduncular nuclei) and parabrachial nucleus. Scattered single projections are obtained from all hypothalamic parts (most of all the ventromedial nucleus), reticular nuclei of the septum, substantia innominata, substantia nigra, truncal nuclei of the raphe. Variety of the dorsal thalamic nuclei, sending their fibers to the BLAC reflects variety of sensory information, that gets here, according to its modality, degree of its differentiation and integrity. A number of the dorsal thalamus nuclei, owing to abundance of labelled neurons, can be considered as special relay thalamic nuclei for the BLAC resembling corresponding relay nuclei for the new cortex.     

A crossed projection from the optic tectum to craniocervical premotor areas in the brainstem reticular formation. An anterograde and retrograde tracing study in the mallard (Anas platyrhynchos L.)

The optic tectum in birds receives visual information from the contralateral retina. This information is passed through to other brain areas via the deep layers of the optic tectum. In the present study the crossed tectobulbar pathway is described in detail. This pathway forms the connection between the optic tectum and the premotor area of craniocervical muscles in the contralateral paramedian reticular formation. It originates predominantly from neurons in the ventromedial part of stratum griseum centrale and to a lesser extent from stratum album centrale. The fibers leave the tectum as a horizontal fiber bundle, and cross the midline through the caudal radix oculomotorius and rostral nucleus oculomotorius. On the contralateral side fibers turn to ventral and descend caudally in the contralateral paramedian reticular formation to the level of the obex. Labeled terminals are found in the ipsilateral medial mesencephalic reticular formation lateral to the radix and motor nucleus of the oculomotor nerve, and in the contralateral paramedian reticular formation, along the descending tract. Neurons in the medial mesencephalic reticular formation in turn project to the paramedian reticular formation. Through the crossed tectobulbar pathway visual information can influence the activity of craniocervical muscles via reticular premotor neurons.     

Efferent connections of the centrum medianum of the cat thalamus revealed by autoradiography

Efferent connections of the centrum medianum and parafascicular nucleus of the thalamus (CM-Pf complex) in cats were studied by the method of anterograde axonal transport of tritiated amino acids followed by autoradiography. Projections from CM-Pf ascend to nuclei of the ventral group and nonspecific nuclei of the thalamus, preoptic, dorsal, lateral, and posterior areas of the hypothalamus, and also into the subthalamic region. Descending pathways are formed only by neurons of the caudomedial part of CM-Pf. They project into the pretectal region, superior colliculus, reticular formation, locus coeruleus, region of the ramus communicans, and substantia grisea centralis of the mesencephalon and pons, and also into the nuclei raphe, magnocellular reticular area, and inferior olivary nucleus of the medulla. In agreement with previous observations it was found that the caudomedial part of CM-Pf does not send direct projections into the cortex and striatum.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 16, No. 2, pp. 224–230, March–April, 1984.     

Neuronal reactions in the reticular nucleus of the thalamus upon realization of conditioned placing and its differential inhibition

Cats were trained to perform conditioned forepaw placing (CFP) using food reinforcement. Reactions of 33 neurons were recorded extracellularly in the reticular nucleus (RN) of the thalamus; among these neurons, 24 units were localized in the sensorimotor part of the nucleus. We used short acoustic signals as conditioning positive and differential (negative) stimuli (PS and DS, respectively). When CFP was realized, 21 neurons of the RN sensorimotor part generated late long-lasting activating or inhibitory spike reactions (durations up to 6 sec and 2 sec, respectively); in 3 neurons there were no reactions. Under conditions of differential inhibition, such reactions were not observed, while the background impulse activity became clearly suppressed. In two-thirds of the neurons under study, alternating PS and DS presentations led to suppression of the reactions to PS presentation. We suppose that differential inhibition-related suppression of the background activity and of the spike reactions related to CFP realization result from restriction of the input synaptic volleys to the RN neurons.     

Vestibulofugal projections to the lateral reticular nucleus of cat medulla

Vestibular nucleus neurons projecting to the cat bulbar lateral reticular nucleus were revealed using horseradish peroxidase axonal transport techniques. Neurons giving rise to such projections — relatively few in number — were confined to homolateral locations and nearly all occurred within Deiter's nucleus. Large as well as small and medium-sized neurons of the vestibular nucleus projected to the lateral reticular nucleus. The part played by the vestibuloreticular projections under review in the control of motor activity is discussed.Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 21, No. 2, pp. 147–152, March–April, 1989.     

Role of the reticular formation of the midbrain in the process of formation of background activity of cortex neurons

In strict experimental conditions the basic activity of optic-cortex neurons in the rabbit was depressed after transverse section at the level of the rostral part of the reticular formation of the midbrain. Electrolytic destruction or functional blockage of the midbrain reticular formation (nucleus reticularis tegmenti) produces a decrease in frequency and magnitude in the grouping indexes of the cortex-neuron pulses that manifest the activity in these conditions. Destruction of specific nuclei in the optic pathway (those of the lateral geniculate body and the corpora bigemina) made no substantial change in the nature of the cortex-neuron pulses. Comparison of the parameters of pulsation activity of the same cortex neurons, as recorded before and after functional exclusion of the midbrain reticular formation, revealed that the increase in grouping of these pulses after the reticular formation was blocked induced changes in the intervals between groups of pulses, while the frequency of pulses within the groups remained constant. On the basis of the data obtained we may assume that the midbrain reticular formation plays an important role in generation of the background activity of cortex neurons, being a triggering mechanism that sets off a group of pulses. Distribution of pulses within the group is apparently due to the activity of cortex mechanisms only.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 2, No. 1, pp. 43–51, January–February, 1970.     

Differential effect of antiepileptic and non-antiepileptic drugs on the reticular formation

The effect of the antiepileptic drugs carbamazepine and phenytoin, and of the non-antiepileptic drug baclofen, was compared on various inhibitory and excitatory mechanisms in the feline trigeminal nucleus. Baclofen resembled carbamazepine and phenytoin in depressing segmental excitatory and facilitating segmental inhibitory mechanisms. However, baclofen facilitated the periventricular and periaqueductal inhibition of the trigeminal nucleus, while carbamazepine and phenytoin depressed these descending inhibitory mechanisms. Baclofen also resembles carbamazepine and phenytoin in its effectiveness in trigeminal neuralgia, but baclofen is not a clinically effective antiepileptic agent. Our experiments indicate that the ability to depress the reticular formation of the diencephalon and midbrain is an important characteristic of antiepileptic drugs. This suggests that the reticular core is involved in the spread and generalization of seizures.     

The organization of the reptilian brainstem reticular formation: A comparative study using Nissl and Golgi techniques

The brainstem reticular formation has been studied in 16 genera representing 11 families of reptiles. Measurements of Nissl-stained reticular neurons revealed that they are distributed along a continuum, ranging in length from 10 μm to 95 μm. Reticular neurons in crocodilians and snakes tend to be larger than those found in lizards and turtles. Golgi studies revealed that reticular neurons posess long, rectilinear, sparsely branching dendrites. Small reticular neurons ( > 31 μm length) possess fusiform or triangular somata which bear two or three primary dendrites. These dendrites have a somewhat simpler ramification pattern when compared with those of large reticular neurons (< 30 μm length). Large reticular neurons generally possess perikarya which are triangular or polygonal in shape. The somata of large reticular neurons bear an average of four primary dendrites. The dendrites of reptilian reticular neurons ramify predominantly in the transverse plane and are devoid of spines or excrescences. The dendritic ramification patterns observed in the various repitilian reticular nuclei were correlated with known input and output connections of these nuclei. Nissl and Golgi techniques were used to divide the reticular formation into seven nuclei. A nucleus reticularis inferior (RI) is found in the myelencephalon, a reticularis medius (RM) in the caudal two-thirds of the metencephalon, and a reticularis superior (RS) in the rostral metencephalon and caudal mesencephalon. Reticularis inferior can be subdivided into a dorsal portion (RID) and a ventral portion (RIV). All reptilian groups possess RID and RM but RIV is lacking in turtles. Reticularis superior can be subdivided into a large-celled lateral portion (RSL) and a small-celled medial portion (RSM). All reptilian groups possess RSM and RSL, but RSL is quite variable in appearance, being best developed in snakes and crocodilians. The myelencephalic raphe nucleus is also quite variable in its morphology among the different reptilian families. A seventh reticular nucleus, reticularis ventrolateralis (RVL), is found only in snakes and in teiid lizards. It was noted that the reticular formation is simpler (fewer numbers of nuclei) in the representatives of older reptilian lineages and more complex (greater numbers of nuclei) in the more modern lineages. Certain reticular nuclei are present or more extensive in those families which have prominent axial musculature.