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.     

Corticalization of two divisions of the visual system during vertebrate evolution

Data on the evolution of the visual system in vertebrate phylogeny are described. Visual projections are demonstrated in the telencephalon of cyclostomata (lampreys). The existence of a retino-thalamo-telencephalic pathway is demonstrated in elasmobranchs (skates). Two visual pathways are present in amphibians (frogs) and reptiles (turtles): retino-thalamo-telencephalic and retino-tecto-thalamo-telencephalic, and these overlap partly at the thalamic level in the lateral geniculate nucleus and completely in the telencephalon. In turtles the earliest visual and tectal impulses relay on their way to the telencephalon in the lateral geniculate body, and later impulses relay in the nucleus rotundus. In mammals (rats) visual tecto-cortical connections are seen; judging from the latent period of potentials arising in the visual cortex in response to stimulation of the superior colliculi these connections have one synaptic relay in the thalamus. The much shorter latent periods of visual evoked potentials recorded in the tectum of the monkey than in turtles (under identical chronic experimental conditions) confirm the views of morphologists on the progressive development of the tectal division of the visual system in vertebrate phylogeny. It is concluded that corticalization of both divisions of the visual system, i.e., the existence of telencephalic representation, appears in the early stages of vertebrate evolution.     

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.     

THE CENTRAL NERVOUS SYSTEM OF CHAMELEON VULGARIS

An analysis of the brain of Chameleon has given evidence of a number of modifications in the general plan of the reptilian nervous system. The telencephalon shows a reduced olfactory mechanism and suggests the presence of the true motor cortex. The ventral peduncle of the lateral forebrain bundle is unusually large with the majority of its fibres terminating in two new nuclear masses in the hypothalamus, the nucleus entopeduncularis, and a few in the nucleus ruber. A high degree of effective response to optic stimuli is indicated by the extensive correlation between the optic tectum and the nuclear masses of the midbrain and the diencephalon. The wide range of eye movements is probably correlated with the development of the basal optic root ganglion and its numerous secondary connections and the elaborate relationship of the geniculate complex. The usually highly developed tecto-bulbar system is partly replaced by a large tegmento-bulbar system that relays the tectal impulses caudally. An interesting mesencephalic nucleus is a large nucleus profundus mesencephali that receives a great variety of impulses and appears to be an important correlation center. Ascending visceral mechanisms were described bringing the hypothalamus into relationship with the lower centers. The great excursion of the tongue is associated with a highly differentiated hypoglossal nucleus. Components of the medial lemniscus from the spinal cord and the medulla to the vicinity of the thalamus were demonstrated.     

Evoked potentials to electrical stimulation of the facial nerve in the carp tectum mesencephali

Tectal evoked potentials to stimulation of the facial nerve, containing afferent fibers of nonolfactory chemoreception, in the carp are positive evoked potentials with a latent period of 5 to 25 msec which show no phase shift as the microelectrode is advanced to a depth of 600 µ. Depending on the amplitude and latency of evoked potentials seven active zones differing in one or both parameters were distinguished in the ipsilateral tectum mesencephali. The role of impulses from the medulla in the mechanism of tectal evoked potentials to facial nerve stimulation is proved by differences in latent periods and disappearance of the tectal response (although it is preserved in the primary center) after severance of connections between the two parts of the brain. Descending influences from the tectum on the primary center were found: its extirpation disturbs evoked potential generation in several parts of the medullla, so that they either disappear completely or their parameters are modified.A. A. Zhdanov State University, Leningrad. Translated from Neirofiziologiya, Vol. 8, No. 1, pp. 39–46, January–February, 1976.     

Visual projections in the forebrain of the carp Cyprinus carpio]

In the carp C. carpio, studies have been made on the evoked potentials (EP) in the-forebrain elicited by electrical stimulation of contralateral (with respect to the midbrain tectum) nerve. The most high-amplitude and short-latency EP were recorded in the rostro-lateral part of the forebrain at the depth of 1,000-1,500 mu. According to the topographical nomenclature of cyprinid telencephalon, this region corresponds to the area dorsalis telencephali pars lateralis, whereas according to classic comparative neuroanatomical nomenclature it may be defined as the primordium hippocamp or primordium piriform zone. Comparison of the EP in the telencephalon with those in the midbrain tectum suggests active nature of the former.     

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).     

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.     

Patterns of cellular proliferation and migration in the developing tectum mesencephali of the frog Rana temporaria and the salamander Pleurodeles waltl

The development of the tectum mesencephali was studied in the frog Rana temporaria and the salamander Pleurodeles waltl by means of nuclear staining and by labeling of cells with bromodeoxyuridine (BrdU). The general spatial and temporal pattern of cell proliferation and cell migration is the same in both species, despite drastic differences in overall tectal morphology. However, the salamander species differs from the frog species by (1) a generally lower cell proliferation rate, (2) a reduction in the activity of the lateral proliferation zone, and (3) a reduction in the formation of superficial cellular layers. Because point (3) affects processes that occur late in ontogeny, our experiments provide evidence that the simple morphology of the tectum of Pleurodeles waltl, compared with the multilayered tectum of Rana, is a consequence of a paedomorphic alteration of the ancestral developmental pattern of the amphibian tectum.     

Tecto-thalamo-telencephalic interrelationships following visual deafferentation

21-23 months after uni- and bilateral enucleation of the eyes, the turtles Emys orbicularis and Testudo horsfieldi exhibited atrophy, gliosis and residual degenerative changes in the visual nerves and tracts and in the two upper layers of the optic tectum, as well as a reduced density of fibres in the tecto-thalamic tract on the deafferentated side. Electrophysiological experiments on the turtles have shown that tectal impulses en route to the general cortex of the forebrain are relayed in the n. rotundus. Conduction of impulses along the tecto-geniculo-cortical path, found in intact animals, ceases, which is apparently due to transsynaptic changes in the surface layers of the optic tectum.     

Hypothalamic unit responses to impulses from the hippocampus and mesencephalic reticular formation

The effect of stimulation of the dorsal tegmentum mesencephali and dorsal hippocampus on spontaneous single unit activity of the retrochiasmal (RCA), lateral (LHA), medial dorsal (MHAd) and medial ventral (MHAv) hypothalamic regions was investigated in acute experiments on rabbits. During repetitive mesencephalic stimulation many more neurons were activated in all parts of the hypothalamus than during hippocampal stimulation. During mesencephalic stimulation more neurons were excited than inhibited in LHA, MHAd, and MHAv, but in RCA more were inhibited. During hippocampal stimulation spontaneous activity was inhibited in many more cells than during mesencephalic stimulation. Neurons on which convergence of impulses from the hippocampus and mesencephalon was observed numbered 25% in RCA, 39% in LHA, 46% in MHAd, and 29% in MHAv of the total number recorded. By their action (inhibitory or excitatory) on the hypothalamic neurons the mesencephalon and hippocampus were antagonistic in 60% of cases in RCA, 78% in LHA, and 61% in MHAd.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 6, No. 5, pp. 489–495, September–October, 1974.     

Postsynaptic inhibition in the primordial hippocampus of the frog

Postsynaptic responses of neurons of the primordial hippocampus to electrical stimulation of brain structures belonging to the visual and olfactory afferent systems were investigated in frogs (Rana temporaria) immobilized with diplacin. * Short-latency (early) and long-latency (late) IPSPs evoked by both olfactory and visual afferent inputs, most probably activated by different conducting systems, are described. Impulses of different modalities can induce both similar and different IPSPs in a neuron. The conducting systems may have a common interneuron for the visual input and specific interneurons for the olfactory input. IPSPs evoked by visual impulses were similar in location to the early IPSP of the olfactory afferent input. Convergence of the systems of early and late inhibition on one neuron was frequently observed for the olfactory afferent input.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 5, No. 6, pp. 583–592, November–December, 1973.     

Convergence of corticofugal impulses on pontine neurons

Convergence of corticofugal impulses in reticular and intrinsic pontine nuclei during stimulation of the frontobasal cortex (proreal, posterior orbital, and basal temporal regions) and also of the dorsal hippocampus was studied in acute experiments on cats anesthesized with a mixture of pentobarbital and chloralose. Three foci of convergence of corticofugal impulses were found in these structures: one in the reticular formation and two in the intrinsic nuclei—in their medial and lateral portions. Neurons with an excitatory type of response were shown to predominate in the reticular formation and neurons with an inhibitory or mixed type of response of neurons activated antidromically by stimulation of one brain region and synaptically during stimulation of another, that the pontine nuclei play an integrative role in the functional unification of structures of the frontobasal zones of the neocortex and hippocampus.M. Gor'kii Donetsk Medical Institute. Translated from Neirofiziologiya, Vol. 12, No. 5, pp. 472–480, September–October, 1980.     

Tegmento-telencephalic relations in turtles

Electrophysiological characteristics of tegmental projections in the forebrain were studied in immobilized turtles (Emys orbicularis), anesthetized with chloralose. The main zone of representation of the tegmentum is located in the striatum, where evoked responses with shorter latencies were recorded and a larger number of units responding distinctly to tegmental stimulation was concentrated than in the dorsal ventricular ridge and general cortex. At the striatal level predominance of lateral tegmental projections was found in the lateral zone and of medial tegmental projections in the medial zone of the striatum. An inhibitory action of conditioning stimulation of the tegmentum on somatic evoked potentials and single unit responses was established. An attempt was made to compare the tegmento-telencephalic system in reptiles and mammals.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 12, No. 3, pp. 255–263, May–June, 1980.     

Homotopic and heterotopic transplantations of quail tectal primordia in chick embryos: Organization of the retinotectal projections in the chimeric embryos

To study the adaptative capabilities of the retinotectal system in birds, the primordium of one optic tectum from 12-somite embryos of Japanese quail was transplanted either homotopically, to replace the ablated same primordium, or heterotopically, to replace the ablated dorsal diencephalon in White Leghorn chick embryos of the same stage. The quail nucleolar marker was used to recognize the transplants. The cytoarchitecture of the tecta and the retinal projections from the eye contralateral to the graft were studied on the 17th or 18th day of incubation in the chimeric embryos by autoradiographic or horseradish peroxidase tracing methods. Morphometric analysis was applied to evaluate the percentage of the tectal surface receiving optic projections. It was observed that: (i) quail mesencephalic alar plate can develop a fully laminated optic tectum even when transplanted heterotopically; (ii) retinal ganglion cells from the chick not only recognize the tectal neurons of the quail as their specific targets in homotopic grafts, but the optic fibers deviate to innervate the heterotopically grafted tectum; (iii) in the presence of a graft, the chick retina is unable to innervate a tectal surface of similar or larger size than that of the control tectum; (iv) tectal regions devoid of optic projections, whether formed by donor or by host cells, always present an atrophic lamination; (v) the diencephalic supernumerary optic tectum competes with and prevails over the host tectum as a target for optic fiber terminals.     

The lateral line mechanoreceptive mesencephalic,diencephalic, and telencephalic regions in the thornback ray,Platyrhinoidis triseriata (Elasmobranchii)

Central lateral line pathways were mapped in the thronback ray, Platyrhinoidis triseriata, by analyzing depth profiles of averaged evoked potentials (AEPs), multiunit activity (MUA), and single unit recordings. Neural activity evoked by contra- or ipsilateral posterior lateral line nerve (pLLN) shock is restricted to the tectum mesencephali, the dorsomedial nucleus (DMN) and anterior nucleus (AN) of the mesencephalic nuclear complex, the posterior central thalamic nucleus (PCT), the lateral tuberal nucleus of the hypothalamus, and the deep medial pallium of the telencephalon (Figs. 2, 3, 4, 6, 7). Neural responses (AEPs and MUA) recorded in different lateral line areas differ with respect to shape, dynamic response properties, and/or latencies (Figs. 9, 10 and Table 1). Ipsilaterally recorded mesencephalic and diencephalic AEPs are less pronounced and of longer latency than their contralateral counterpart (Fig. 9 and Table 1). In contrast, AEP recorded in the telencephalon show a weak ipsilateral preference. If stimulated with a low amplitude water wave most DMN, AN, and tectal lateral line units respond in the frequency range 6.5 Hz to 200 Hz. Best frequencies (in terms of least displacement) are 75-150 Hz with a peak-to-peak water displacement of 0.04 micron sufficient to evoke a response in the most sensitive units (Fig. 11A, B, C). DMN and AN lateral line units have small excitatory receptive fields (RFs). Anterior, middle, and posterior body surfaces map onto the rostral, middle, and posterior brain surfaces of the contralateral DMN (Fig. 12). Some units recorded in the PCT are bimodal; they respond to a hydrodynamic flow field--generated with a ruler approaching the fish--only if the light is on and the eye facing the ruler is left uncovered (Fig. 13).     

Cellular migration and morphological complexity in the caecilian brain

The morphology of the tectum mesencephali and the medial pallium is studied in species representing the six families of caecilians (Amphibia: Gymnophiona) in order to determine whether differences in brain morphology are related to function, phylogenetic history, or life history strategies. In general, the caecilian tectum is characterized by simplification in having little to no lamination and few migrated cells. The degree of morphological complexity differs between species and between brain regions. Our data suggest that changes in brain morphology are due to a mosaic of different influences. We did not find a strict correlation between visual system reduction and tectal morphologies. However, phylogenetic effects exist. The greatest degree of morphological complexity is found in members of the Rhinatrematidae, a family that is considered basal to the lineage. Thus, simplification of brain morphology in caecilians must be considered a secondary or derived rather than a primitive feature. Direct development and miniaturization are correlated with the greatest simplification in the tectum mesencephali and medial pallium. There is a relationship between differences in brain morphology and heterochrony in caecilians, as in other amphibians. J. Morphol. 231:11–27, 1997. © 1997 Wiley-Liss, Inc.     

Visual Stimuli Evoked Action Potentials Trigger Rapidly Propagating Dendritic Calcium Transients in the Frog Optic Tectum Layer 6 Neurons

The superior colliculus in mammals or the optic tectum in amphibians is a major visual information processing center responsible for generation of orientating responses such as saccades in monkeys or prey catching avoidance behavior in frogs. The conserved structure function of the superior colliculus the optic tectum across distant species such as frogs, birds monkeys permits to draw rather general conclusions after studying a single species. We chose the frog optic tectum because we are able to perform whole-cell voltage-clamp recordings fluorescence imaging of tectal neurons while they respond to a visual stimulus. In the optic tectum of amphibians most visual information is processed by pear-shaped neurons possessing long dendritic branches, which receive the majority of synapses originating from the retinal ganglion cells. Since the first step of the retinal input integration is performed on these dendrites, it is important to know whether this integration is enhanced by active dendritic properties. We demonstrate that rapid calcium transients coinciding with the visual stimulus evoked action potentials in the somatic recordings can be readily detected up to the fine branches of these dendrites. These transients were blocked by calcium channel blockers nifedipine CdCl₂ indicating that calcium entered dendrites via voltage-activated L-type calcium channels. The high speed of calcium transient propagation, >300 μm in <10 ms, is consistent with the notion that action potentials, actively propagating along dendrites, open voltage-gated L-type calcium channels causing rapid calcium concentration transients in the dendrites. We conclude that such activation by somatic action potentials of the dendritic voltage gated calcium channels in the close vicinity to the synapses formed by axons of the retinal ganglion cells may facilitate visual information processing in the principal neurons of the frog optic tectum.     

Rotation of the tectal primordium reveals plasticity of target recognition in retinotectal projection

Retinotectal projection is precisely organized in a retinotopic manner. In normal projection, temporal retinal axons project to the rostral part of the tectum, and nasal axons to the caudal part of the tectum. The two-dimensional relationship between the retina and the tectum offers a useful experimental system for analysis of neuronal target recognition. We carried out rotation of the tectal primordium in birds at an early stage of development, around the 10-somite stage, to achieve a better understanding of the characteristics of target recognition, especially the rostrocaudal specificity of the tectum. Our results showed that temporal retinal axons projected to the rostral part of the rotated tectum, which was originally caudal, and that nasal axons projected to the caudal part of the rotated tectum, which was originally rostral. Therefore, the tectum that had been rotated at the 10-somite stage received normal topographic projection from the retinal ganglion cells. Rostrocaudal specificity of the tectum for target recognition is not determined by the 10-somite stage and is acquired through interactions between the tectal primordium and its surrounding structures.