The tecto-thalamo-telencephalic system of the tortoise (electrophysiological investigation)

Using tortoises immobilized with diplacin to which chloralose had been added, or under chloralose — nembutal narcosis, we investigated the distribution of evoked potentials and neuronal responses in the thalamus and forebrain induced by electrical excitation of the optic tectum and by flashes of light. In the thalamus the main mass of cells that reacted to these stimuli was concentrated in the nucleus rotundus and the tecto-thalamic tract; in the forebrain it was concentrated in the general cortex, the pallial thickening, and the neostriatum. In the two latter structures responses with shorter latent periods than those in the general cortex predominated. Visual and tectal neuronal responses, especially those of convergent cells, displayed correlation in their latent periods and types of response, which was more clearly shown in the thalamus. Submaximal tetanization of the optic tectum had a facilitating effect on cortical responses produced by light flashes and excitation of the nucleus rotundus. Complete blocking of transmission of tectal impulsation to the forebrain was observed on destruction of the tecto-thalamic tract region bounded by the lateral bundle of the forebrain, the lateral geniculate body, and the nucleus rotundus. High-frequency excitation of the nucleus rotundus produced only partial blocking of transmission (of the late components). It is concluded that there are various pathways of tectal impulsation through the thalamus to the forebrain.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 2, No. 3, pp. 296–306, May–June, 1970.     

Electrophysiological analysis of cortico-tectal connections in the turtle

Two types of evoked potentials are recorded in the tectum mesencephali in response to electrical stimulation of the forebrain surface of the turtleEmys orbicularis. The results of a layer-by-layer analysis show that evoked potentials of type I in response to stimulation of the hippocampal and piriform cortex are generated outside the tectum. Evoked potentials of type II, consisting of two surface-negative components, are recorded in the tectum in response to stimulation of the rostro-central surface of the forebrain. The first component appeared after a latent period of 20 msec and lasted 40–60 msec; the second component appeared after 80–100 msec and lasted 100–300 msec. Layer-by-layer and pharmacological analysis showed that the first component of the type II evoked potential is generated in the tegmental structures of the mesencephalon, whereas the second (long-latency) is generated in the tectum. The tectal origin of the second component is confirmed by its interaction with the tectal response to photic stimulation or to electrical stimulation of the optic nerve, evidence that these evoked potentials are generated by common structures. The efferent pathway from the dorsal cortex to the primary visual center is unilateral and has features of polysynaptic projections (long latent period, low lability).     

Direct cortical response in the cat motor cortex during summation

Summation was studied by a procedure close to that used in producing a conditioned reflex. Subthreshold electrical stimulation, which gave rise to a dominant focus in the cat motor cortex, was applied after photic stimulation. Under these conditions, summation occurred both when the two stimuli were applied simultaneously and when the weaker stimulus preceded the stronger one by a very short interval (tens of milliseconds). Increased excitability was characteristic of the dominant focus. An excessive increase in excitability weakened the summation reflex. Electrographically, this type of conditioning was reflected in an increase in amplitude of the primary negative wave of the direct cortical response, recorded in the motor area at a distance of 2–3 mm from the stimulation point. It is concluded from analysis of this electrophysiological phenomenon and comparison of the results with data in the literature that different mechanisms are involved in the summation process during different sequences of stimulation ("photic+electrical" and "electrical+photic").Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 1, No. 3, pp. 293–302, November–December, 1969.     

Early positive component of the tectal evoked potential ofSqualus acanthias during electrical stimulation of the optical nerve

The evoked potential of the tectum opticum during electrical stimulation of the optic nerve was studied in acute experiments on the dogfishSqualus acanthias L. The negative phase of the "classical" negative-positive evoked potential of the contralateral hemisphere of the tectum opticum was shown to be a complex potential, including an early positive component. A similar potential also was recorded from the ipsilateral hemisphere. Enhancement of this positive potential on insertion of the recording electrode deep into the brain, its resistance to functional block on application of potassium chloride to the brain surface, and recording a similar potential from the surface of the floor of the third ventricle after extirpation of the tectum opticum are evidence of the nontectal location of the source of this evoked potential component. On the basis of existence of a focus of maximal activity in the rostral zones of the brain beneath the tectum opticum, and disappearance of the early positive component during functional block and extirpation of this brain region, it is concluded that a leading role in the generation of this component is played by thalamic nuclei.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 16, No. 1, pp. 61–67, January–February, 1984.     

Functional and ultrastructural changes in the midbrain tectum of the frog after enucleation

In experiments on frogs immobilized with Diplacin at different times after unilateral enucleation, as degeneration of the optic fibers and their terminals developed, a successive disappearance of the components of evoked potential (EP) was observed; this indicates the direct dependence of the rate of degeneration on the diameters of the fibers. The nature of the ultrastructural changes also depends on the diameter of the cut axons: the terminals of all or some of the myelinated fibers of large diameter degenerated with the condensation of the endings and of all the cytoplasmatic organelles ("dark" type); the terminals of thin myelinated and unmyelinated fibers degenerated with a gradual lysis of organelles ("light" type). Unmyelinated optic fibers and their synapses survived and transmitted the excitation for more than 140 days at a temperature of 18–20°C. In the course of the survival of optic fibers and synapses, the static (latency, duration, and amplitude) as well as dynamic (lability, excitability cycle, and posttetanic changes) characteristics of the EP for electrical stimulation of the nerve changed insignificantly. Direct response of the midbrain tectum decreased in the course of the degeneration of the optic fibers, and after 280 days its amplitude was about 20% of the control value.A. N. Severtsov Institute of Evolutionary Morphology and Ecology of Animals, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 2, No. 2, pp. 180–188, March–April, 1970.     

Excitation and inhibition in the visual center of Rana temporaria

Analysis of postsynaptic unit responses in the visual center ofRana temporaria showed that optic nerve fibers with high and low conduction velocities usually converge on a single neuron of the tectum opticum (TO). In response to stimulation of the optic nerve a complex depolarization potential consisting of 3 (or possibly 4) EPSPs was recorded in one group of neurons; these EPSPs were probably generated through excitation of several groups of afferent fibers. Either an increase or a decrease in the EPSPs can be observed in the TO neurons in response to repetitive and paired stimulation of the optic nerve. Postsynaptic inhibitory responses of some TO neurons, probably of direct and recurrent origin, are discussed.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 3, No. 6, pp. 637–643, November–December, 1971.     

Response of neostriatal neurons to direct electrical stimulation of the optic tract and photic stimulation in awake cats

During acute experiments on awake cats the response of 98 neurons belonging to the head and tail of the caudate nucleus to direct electrical stimulation of the optic tract and presentation of photic stimuli was investigated using extracellular recording techniques. Of the test neurons 34.6% responded to stimulation of the optic tract and 36.2% to optic stimulation. Long latency (over 40 msec for the optic tract and over 80 msec for visual stimulation) excitatory responses prevailed in both cases. A small number of cells responded to optic tract stimulation with short latencies of 5–14 msec. Both types of stimulation were presented during investigations of 58 units of which eight were found to respond to both stimuli. The latter varied in their reaction to different stimuli and their response pattern. Findings are discussed in relation to the possible pathways by which visual information reaches the cortical structure under study.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 18, No. 4, pp. 476–485, July–August, 1986.     

Electrical potentials from the eye and optic nerve of Strombus: effects of electrical stimulation of the optic nerve.

1. Photic stimulation of the mature eye of Strombus can evoke in the optic nerve 'on' activity in numerous small afferent fibres and repetitive 'off' bursts of afferent impulses in a smaller number of larger fibres. 2. Synchronous invasion of the eye by electrically evoked impulses in small optic nerve fibres (apparently the 'on' afferents, antidromically activated) can evoke a burst of impulses in the larger 'off' fibres which propagate away from the eye. Invasion of the eye via one branch of optic nerve can evoke an answering burst in another branch. 3. Such electrically evoked bursts are similar to light-evoked 'off' bursts with respect to their impulse composition, their ability to be inhibited by illumination of the eye, and their susceptibility to MgCl2 anaesthesia. 4. Invasion of the eye by a train of repetitive electrically evoked impulses in the absence of photic stimulation can give rise to repetitive 'off' bursts as well as concomitant oscillatory potentials in the eye which are similar to those normally evoked by cessation of a photic stimulus. 5. The electrically evoked 'off' bursts appear to be caused by an excitatory rebound following the cessation of inhibitory synaptic input from photoreceptors which can be antidromically activated by electrical stimulation of the optic nerve. 6. The experimental results suggest that the rhythmic discharge of the 'off' fibres evoked by the cessation of a photic stimulus is mediated by the abrupt decrease of inhibitory synaptic input from the receptors.     

Nonolfactory afferent telencephalic projections in frogs

The degree of modal specificity of neurons of the tectum, dorsal thalamus, and primordial hippocampus relative to visual (photic) and somatic (tactile) stimulation was investigated in experiments on frogs (Rana temporaria) immobilized with tubocurarine. The specificity of sensory information arriving at the successive levels of the frog's thalamo-telencephalic afferent system diminishes in an ascending direction. Evoked potentials arising in the telencephalon in response to electrical stimulation of the sciatic nerve and to flashes are found not only in the primordial hippocampus, but also bilaterally in the septum and striatum. It is concluded on the basis of discharges recorded in the fibers of the thalamo-telencephalic tracts during somatic and visual stimulation that fibers carrying nonolfactory information into the telencephalon run in the medial and lateral forebrain bundles.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. International Brain Research Laboratory, Kotor, Yugoslavia. Translated from Neirofiziologiya, Vol. 5, No. 5, pp. 537–544, September–October, 1973.     

Responses of neurons of reticular and ventral anterior nuclei of the cat thalamus to afferent stimuli of different modalities

Responses of 146 spontaneously active neurons of the reticular nucleus (R) and of 98 neurons of the ventral anterior (VA) nucleus of the thalamus to electrical stimulation of the skin of the footpads, to flashes, and to clicks were studied in experiments on cats immobilized with D-tubocurarine or myorelaxin. Stimulation of the contralateral forelimb was the most effective: 24.9% of R neurons and 31.3% of VA neurons responded to this stimulation. A response to clicks was observed in only 4.4% of R neurons and 2.4% of VA neurons. Nearly all responding neurons did so by phasic (one spike or a group of spikes) or tonic excitation. Depression of spontaneous activity was observed only in response to electrical stimulation of the skin. Depending on the site of stimulation, it was observed in 2.6–4.3% of R neurons and 1.7–2.1% of VA neurons tested. The latent period of the phasic responses of most neurons was 6–64 msec to electrical stimulation of the contralateral forelimb, 11–43 msec in response to stimulation of the hindlimb on the same side, 10–60 msec to photic and 8–60 msec to acoustic stimulation. Depending on the character of stimulation, 75.1–95.6% of R neurons and 68.7–97.6% of VA cells did not respond at all to the stimuli used. Of the total number of cells tested against the whole range of stimuli, 25% of R neurons and 47% of VA neurons responded to stimulation of different limbs, whereas 16% of R neurons and 22% of VA cells responded to stimuli of different sensory modalities. The functional role of the convergence revealed in these experiments is to inhibit (or, less frequently, to facilitate) the response of a neuron to a testing stimulus during the 40–70 msec after conditioning stimulation.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 7, No. 6, pp. 563–571, November–December, 1975.     

Restoration of retino-tectal connections in frogs during regeneration of the optic nerve

At various times after unilateral division of the optic nerve in the frogRana temporaria L. evoked potentials in response to electrical stimulation of the optic nerve were investigated in a segment distal to the site of operation, spike activity was recorded from endings of regenerating and intertectal axons when stimuli of different shapes were placed in the field of vision, and the distribution of axonal bulbs of growth by depth in the tectum mesencephal was studied electron-microscopically. During regeneration of the axons the responses of the retinal ganglionic cells to visual stimuli retained most of their individual features. Myelinated axons of the retinal ganglionic cells regenerate first (starting on the 21st day after operation). Myelination of these fibers lags significantly behind their growth and is complete more than 100 days after the operation. Unmyelinated axons of the retinal ganglionic cells grow up toward the tectum mesencephali after myelinated axons (80 or more days after the operation). Axonal bulbs of growth in the initial periods after the operation are located close to the pial surface and the level of spread of the myelinated axons of the retinal ganglionic cells differs significantly from their normal level of localization. Intertectal connections persist after division of the nerve and are activated by visual stimuli during regeneration of the axons of the retinal ganglionic cells. Connections were found mainly between intertectal fibers terminating superficially and retinal ganglionic cells belonging to class 1 and 2 detectors. Axons of the retinal ganglionic cells grow up toward the caudal region of the tectum mesencephali later than toward the rostral region.A. N. Severtsov Institute of Evolutionary Morphology and Ecology of Animals, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 5, No. 6, pp. 611–620, November–December, 1973.     

Transthalamic conduction of visual impulses to the cortex and subcortical forebrain structures in turtles

The thalamic relays for the conduction of impulses arising during photic stimulation of the eyes and electrical stimulation of the tectum in the general cortex, hyperstriatum (the dorsal ventricular ridge), and the striatum proper were studied in the turtleEmys orbicularis. Acute experiments on immobilized animals showed that anodal polarization temporarily and destruction of n. rotundus irreversibly suppress the main negative wave of the responses to tectal stimulation and to flashes in the hyperstriatum, whereas the corresponding responses in the general cortex still persist. Polarization and destruction of the lateral thalamic region, including the lateral geniculate body, have the opposite effect: responses in the hyperstriatum to photic and tectal stimulation are virtually unchanged whereas those in the general cortex disappear, except their late components. Preceding single stimulation of the tectum or n. rotundus depresses responses in the hyperstriatum evoked by flashes. However, during stimulation of the lateral thalamic region, combined potentials and single unit responses appear in the hyperstriatum and interact with responses evoked by tectal stimulation. It is concluded that the main pathways in turtles which supply visual information to the general cortex and hyperstriatum differ: the former relay in the lateral thalamic region, the latter in n. rotundus, although some overlapping of their projections in the hyperstriatum and striatum is possible.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Leningrad. Translated from Neirofiziologiya, Vol. 9, No. 5, pp. 486–494, September–October, 1977.     

Functional heterogeneity of lateral geniculate body neurons in the rabbit]

Background and evoked activity of LGB units was studied on immobilized and anaesthetized rabbits. Two groups of projection units were revealed, differing by the level of background activity, latencies and mean frequency of discharges in responses to single photic flashes and to electrical stimulation of the optic nerve. It is assumed that these groups of units belong to the slowly and rapidly conducting paths of sensory information transmission in the visual projection system.     

Responses of neurons of the frog nucleus isthmi to optic nerve stimulation

Responses of single units in the region of the nucleus isthmi were investigated in frogs immobilized with diplacin by means of extracellular capillary liquid microelectrodes. The neurons of this region were not spontaneously active and, after electrical stimulation of the contralateral optic nerve with a single pulse, they gave a single spike discharge with a minimal latent period of 20–110 msec. On increasing the strength of stimulation from threshold to maximal these latent periods were significantly reduced, indicating marked development of summation processes in the corresponding neuronal pathway. Only 14% of the neurons (7 of 57) discharged in response to stimuli exciting myelinated but not exciting unmyelinated optic fibers. All neurons were characterized by instability of the latent period of the evoked discharge; consequently, they were not excited antidromically. The possible functional role of the nucleus isthmi region in the Anura is discussed.A. N. Severtsov Institute of Evolutionary Morphology and Ecology of Animals, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 9, No. 1, pp. 33–40, January–February, 1977.     

Characteristics of dynamic reorganization of receptive fields of cortical and geniculate visual neurons of cat in response to changes in photic stimulation parameters

The effect of parameters of local photic stimulation of different points of the receptive field on the characteristics of dynamic reorganization of receptive fields of cortical and geniculate visual neurons within microintervals of time observed previously was studied in computer-controlled experiments on unanesthetized, curarized cats. Dependence on the degree of widening of the receptive field and the temporal characteristics of this process on the background illumination level, intensity, contrast, area, duration, energy, and orientation of a local rectangular or circular photic stimulus flashing in random order at 100 points of the tested part of the visual field was studied. It was concluded that the background illumination level and the intensity, size, duration, and orientation of the stimulus have a specific effect on dynamic reorganization of the receptive field. The effects of all the parameters studied on the dynamics of the receptive field were shown to be nonlinear functions with optimal values that differed for different cells. The possible functional role of this effect and also the probability that it may participate in information coding in the visual system are discussed.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 15, No. 4, pp. 339–346, July–August, 1983.     

Pretecto-tectal influences

(1)From the dorsal surface of the toad (Bufo b. spinosus, B. marinus) optic tectum (OT), field potentials (FP) were recorded at 9 reference sites in response to electrical stimulation of the optic nerve (ON). The FP showed 4 main components, besides an initial deflection attributed to axonal potentials: two negative waves N1, N2 (attributed to postsynaptic excitatory processes) and two positive waves P2, P3 (attributed to postsynaptic inhibitory processes). The responses across the reference sites were rather similar in different individuals. (2) Electrical stimulation of an area in the ipsilateral pretectal lateral posterodorsal and posterior (Lpd/P) thalamic region evoked tectal FPs showing mainly a negative and a positive wave. Regarding wave amplitudes, the FPs displayed disproportionalities across the reference sites. (3) Electrical stimulation of the contralateral Lpd/P evoked mainly a positive wave in the tectal FP whose disproportionality corresponded roughly to the one obtained to ipsilateral Lpd/P stimulation. (4) The inital negative wave of the tectal FP in response to ON stimulation was nearly abolished, if Lpd/P stimulation preceded ON stimulation at a delay of 17–25 ms. (5) Since FPs showed adaptation to repetitive stimulation, various experiments were carried out to distinguish adaptation phenomena from effects of neuronal interactions between Lpd/P and OT. (6) The results provide evidence that ON- and Lpd/P-mediated inputs interact in superficial tectal layers, whereby pretectotectal input suppresses retinotectal excitatory information transfer. Input of Lpd/P to the contralateral superficial OT suggests postsynaptic inhibition. This study provides no information about pretectal inputs to deeper tectal layers, which anatomically are known to exist.Abbreviations A-I recording sites from the dorsal tectal surface - D _t delay between Lpd/P and ON stimulation - EPSP IPSP excitatory and inhibitory postsynaptic potentials, respectively - FP field potential - L latency of FP waves - ON optic nerve - OT optic tectum - Lpd/P lateral posterodorsal and posterior pretectal thalamic region - Lpv lateral posteroventral pretectal thalamic nucleus - N, P negative and positive waves of FPs, respectively - PRE presynaptic axonal input - TH pretectal thalamic neurons     

Neurophysiological properties of magnetic cells in the pigeon's visual system

Summary Single unit electrical activity was recorded extracellularly in the nucleus of the basal optic root (nBOR) and in the optic tectum under earth-strength magnetic stimulation. Units in the nBOR which were stimulated while the eyes were illuminated by light of different wavelengths exhibited peaks of magnetic responsiveness at 503 nm and 582 nm.Magnetically directional selective cells were found in the stratum griseum et fibrosum superficiale of the optic tectum. They also showed directional selectivity to dynamic photic stimuli. Response peaks varied with the orientation of the pigeon in the horizontal plane. This confirmed that the magnetic responses contained directional information. The results suggest that the receptor and neural organisation of the pigeon's visual system provides an adequate substrate for the detection and elaboration of magnetic compass information.     

Layer by layer analysis of synaptic organization in the optic tectum of the prog

The distribution of axo-axonal and axo-dendritic synapses, nerve endings, and bodies of neurons by depth in the optic tectum ofRana temporaria L. was investigated under normal conditions and 6–9, 60, and 134 days after removal of the contralateral eye. Counting was carried out on long oriented sections examined in the electron microscope. In outer plexiform layer 9 the density of synapses was greatest near the surface of the tectum and decreased in the direction away from it; no inner sublayers with differing density of synapses could be distinguished. In the outer zone of layer 9 (to a depth of about 30 ) many axo-axonal synpases were discovered. Endings of myelinated optic fibers of large diameter ("dark" terminal degeneration) were widely distributed in the same layer. The density of axo-dendritic synapses in deep plexiform layer 5 was similar to that in layer 9. Many nerve endings containing granular vesicles as well as pale synaptic vesicles were found in layer 5 and neighboring zones.A. N. Severtsov Institute of Evolutionary Morphology and Ecology of Animals, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 11, No. 2, pp. 130–136, March–April, 1979.     

Synaptic nature of the principal components of the evoked potential in the general cortex of the turtle forebrain

The nature of the principal components of the evoked potential of the general cortex of the turtle forebrain was studied in response to electrical stimulation of the contralateral optic nerve. Comparison of these components with postsynaptic potentials of the neurons of this structure showed that the four fast negative waves of the evoked potential correspond to fast EPSPs, which are independent of one another. The positive wave of the evoked potential is the sum of several IPSPs. The slow negative and, to some extent, the positive wave are a reflection of the slow EPSP. It is shown that early EPSPs are generated on portions of the apical dendrites which are further from the soma than those generating late fast EPSPs and also the IPSP and slow EPSP. Axo-somatic contacts are perhaps also concerned in the generation of the last-named potential.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 5, No.3, pp.261–271, May–June, 1973.     

Postsynaptic inhibition in the general cortex of the turtle forebrain

Postsynaptic inhibition in the general cortex of the turtle forebrain was investigated by recording unit activity intracellularly. Depending on the type of IPSPs recorded in response to electrical stimulation of the contralateral optic nerve and cortical surface the neurons were subdivided into three groups: 1) with long direct IPSPs, 2) with long and short direct, and also recurrent IPSPs, 3) with short direct and recurrent IPSPs. It is concluded that inhibitory pathways of the short direct and recurrent IPSPs have a common final component, a stellate interneuron. Compared with the recurrent collaterals of the principal neurons, the direct afferents make contact with more distal portions of the dendrites of this cell. Synapses formed on dendrites of the principal neurons by axons of the stellate cells are nearer to the soma than synapses responsible for generation of the long direct IPSP.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 5, No. 4, pp. 375–383, July–August, 1973.