Calcium-Binding Proteins in the Turtle Thalamus. Analysis in the Light of Hypothesis of the “Core-Matrix” Thalamic Organization in Relation to the Problem of Homology of Thalamic Nuclei among Amniotes

Distribution of immunoreactivity (IR) to Ca-binding proteins (CaBPr) (calbindin, Calb, parvalbumin, Parv., and calretinin, Calr) was studied in the thalamus of the Central Asian terrestrial turtles (Testudo horsfieldi) and fresh water turtles (Emys orbicularis). There has been established a wide spread of these proteins, which combines overlapping and a relative alternation of distribution of different CaBPr in individual nuclei. A comparison of IR was made in two relay nuclei of the visual system, GLd and Rot. Both nuclei had IR to all CaBPr, but with different degree of intensity. In the terrestrial turtles, the amounts of Calb-, Parv-, and Calr-IR neurons in the cellular plate of the GLd were close. In this plate and in the neuropil part of this nucleus there was observed CaBPr-innervation of various density. Calr-IR neurons in the GLd of the fresh water turtles dominated over Parv- and Calb-IR neurons, whose detection varied significantly. In Rot, a clear predominance of Calb-IR neurons was shown over Parv- and Calr-IR cells by constancy of their detection, the number (1.5–2-fold higher), and intensity of the immune label, as well as the highest density of Calb and Parv innervation. The character of IR in the Rot was similar in the both turtle species. In the auditory and somatic relay thalamic nuclei and in the non-sensory anterior thalamic nuclei (Dma, Dla) there were present neurons and terminals with IR to all CaBPr without any predominance of Parv-IR in the relay nuclei and Calb-IR in the anterior thalamic nuclei. The constant and characteristic feature of Enta in the turtles of both species is a dense population of Parv-IR neurons, whose topography and cellular composition coincide with those of population of GABA-IR neurons in this nucleus. The data obtained have shown that the alternative presence of different CaBPr in the relay sensory and non-sensory thalamic nuclei, which has been established as a characteristic feature of the mammalian thalamus, is not characteristic at all of turtles. It seems that in the course of evolution there occurred a reorganization of distribution of different CaBPr in thalamic nuclei of amniotes due to changes of their functional loading. The reptilian thalamic sensory relay nuclei are likely to be represented mainly by less specific parts comparable with Calb-IR matrix of specific nuclei in the higher amniotes (mammals), while their more specialized (core) Parv-IR regions are formed later in evolution. Therefore, the distribution of Parv- and Calb-IR neurons in the turtle thalamic nuclei cannot be a criterion at evaluation of homology of thalamic nuclei in amniotes, but permits judging about the degree of their specialization.     

Evolutionary evaluation of reciprocity of connections in the turtle tectofugal visual system

In two turtle species—Emys orbicularis and Testudo horsfieldi—by the method of anterograde and retrograde traicing at the light and electron microscopy level, the existence is proven of direct descending projections from the thalamic nucleus of the tectofugal visual system n. rotunds (Rot) to the optic tectum. After injection of tracers into Rot alone and into Rot with involvement of the tectothalamic tract (Trtth), occasional labeled fibers with varicosities and terminals are revealed predominantly in the deep sublayers of SGFS of the rostral optic tectum, while in the lower amount—in other tectal layers. After the tracer injections into the optic tectum, a few retrogradely labeled neurons were found mainly in the Rot ventral parts and within Trtth. Their localization coincides with that of GABA-immunoreactive cells. Electron microscopy showed the existence of many retrogradely labeled dendrites throughout the whole Rot; a few labeled cell bodies were also present there, some of them being also GABA-immunoreactive. These results allow us to conclude about the existence of reciprocal connections between the optic tectum and Rot in turtles, these connections being able to affect processing of visual information in tectum. We suggest that reciprocity of tectothalamic connections might be the ancestral feature of the vertebrate brain; in the course of amniote evolution the functional significance of this feature can be decreased and even lost in parallel with a rise of the role of direct corticotectal projections.     

Calcium-binding proteins and cytochrome oxidase activity in the turtle optic tectum with special reference to the tectofugal visual pathway

Using immunohistochemistry and a tracer technique we investigated the distribution in the optic tectum of turtles (Emys orbicularis and Testudo horsfieldi) of the calcium-binding proteins (CaBPr) parvalbumin (PV), calbindin (CB) and calretinin (CR) before and after labeling of the nucleus rotundus (Rot) with horseradish peroxidase. The optic tectum activity of the cytochrome oxidase (CO) was studied in parallel. In the principal link of the tectofugal visual pathway (central gray layer, SGC) in both chelonian species, the sparse PV-ir as well as CB- and CR-ir neurons were found significantly varying both in number and the intensity of immunoreactivity of their bodies and dendrites. In contrast, the superficial (SGFS) and deeper periventricular (SGP) tectal layers comprised numerous cells immunoreactive to all three CaBPr in different proportions. Only few retrogradely labeled tectorotundal SGC neurons expressed PV, CB or CR. The very large PV-ir neurons in SGC and SAC were not retrogradely labeled; morphologically they matched the efferent neurons with descending projections. SGC neurons of two chelonian species differed in the level of CO activity. Intense immunoreactivity to all three CaBPr and high CO activity were detected in both species in SGFS neuropil with some differences in sublaminar distribution patterns. The peculiarities of the CaBPr and CO activity distribution patterns in different segments of SGC neurons are discussed as related to the laminar organization of the turtle tectum and its retinal innervation. It is suggested that in the projection tectorotundal SGC neurons the CaBPr are concentrated mainly in their distal dendrites that contact retinal afferents in the superficial retinorecipient tectal layer.     

Activity of cytochrome oxidase in centers of tectofugal and thalamofugal tracts of the visual system of pigeon <Emphasis Type="Italic">Columbia livia</Emphasis>

By using a histochemical method of determination of activity of cytochrome oxidase (CO), the level of metabolic activity in pigeons has been shown to be higher in centers of the tectofugal visual tract (pretectal nuclei: Pr, SP, SP/IPS, thalamic nucleus Rot, end telencephalic entopallidum) than in centers of the thalamofugal visual tract (GLd of the thalamus, visual area of the hyperpallium Wulst). These data agree with the concept of dominating role of the tectofugal visual tract in organization of the bird everyday behavior. The high CO activity is also characteristic of the mesencephalic structures (EM, isthmic nuclei: IMc, IPc, and SLu) modulating transduction of visual information in tectum, Rot, and GLd. Similar differences in the metabolic activities between two visual system tracts were earlier shown in reptiles, which indicates the evolutionary conservatism of the tectofugal visual tract among the sauropside amniotes. However, in pigeons the level of the CO activity in some GLd nuclei approaches that in Rot, which allows us to suggest an increase in birds of role of the thalamofugal tract in processing of information necessary for performance of complex visual functions.     

The level of metabolic activity (cytochrome oxidase) as an index of functional significance of tectofugal and thalamofugal channels of the reptilian visual system

Using histochemical determination of activity of the mitochondrial oxidative enzyme cytochrome oxidase (CO) in brain structures, metabolic activity both in turtles and in lizards has been shown to be higher in centers of the tectofugal channel (the tectal stratum griseum centrale, SGC; nucleus pretectalis ventralis, Ptv; thalamic nucleus rotundus, Rot; telencephalic visual area of the anterior dorsal ventricular ridge, Advr) than in the thalamofugal channel centers (the thalamic nucleus geniculatus lateralis pars dorsalis, GLd; cortex dorsolateralis, Cxdl; and pallial thickening, Path) of the visual system. Some interspecies differences in distribution of the CO activity in the tectal, thalamic, and telencephalic visual centers between terrestrial and pond turtles and lizards were revealed. The obtained data confirm the idea on the dominating role of the tectofugal channel over the thalamofugal channel of the visual system in information processing and organization of the day-to-day behavior of reptiles.     

Activity of cytochrome oxidase in centers of tectofugal and thalamofugal channels of the visual system of pigeon Columba livia

By using a histochemical method of determination of activity of cytochrome oxidase (CO), the level of metabolic activity in pigeons has been shown to be higher in centers of the tectofugal visual channel (pretectal nuclei: Pr, SP, SP/IPS, thalamic nucleus Rot, telencephalic entopallidum) than in centers of the thalamofugal visual channel (GLd, visual area of the hyperpallium Wulst). These data agree with the concept of the dominating role of the tectofugal visual channel in organization of the bird everyday behavior. The high CO activity is also characteristic of the mesencephalic structures (EM, isthmus nuclei: IMc, IPc, SLu) modulating transduction of visual information in tectum, Rot and GLd. Similar differences in the metabolic activities between two visual system channels have been shown earlier in reptiles, which indicates the evolutionary conservatism of the tectofugal visual channel among the sauropside amniotes. However, in pigeons the level of the CO activity in some GLd nuclei approaches that in Rot, which allows us to suggest a rise in birds of the role of the thalamofugal channel in processing of information necessary for performance of complex visual functions.     

Neurons of visual thalamic nuclei projecting to telencephalon express different types of calcium-binding proteins: A combined immunocytochemical and tracer study

In turtles (Testudo horsfieldi, Emys orbicularis), immunoreactivity to calbindin (CB), parvalbumin (PV), calretinin (CR) and co-localization of CB and PV were studied in neurons of the visual thalamic nuclei (Rot, GLd) projecting to the telencephalon using a combination of immunohistochemical and tracer methods. The prevalence of CB-immunoreactive (-ir) neurons in Rot, CB-ir and CR-ir neurons in GLd, and a smaller number of PV-ir neurons in both nuclei was shown. Double immunofluorescent labeling revealed that within both nuclei PV and CB are colocalized in most PV-ir and fewer CB-ir neurons. After injection of horseradish peroxidase into the Rot and GLs telencephalic projection fields, retrograde labeling was found in corresponding thalamic projection neurons immunoreactive to all the three proteins. After introduction of the fluorescent tracer Fluo-gold into the same telencephalic regions, retrograde labeling was detected in Rot and GLd neurons immunoreactive only to PV and CB as well as in neurons with colocalization of both proteins. These findings provide further evidence that in turtles the CB component prevails in the rotundo-telencephalic pathway while the CB/CR component is dominant in the geniculotelencephalic pathway. The role of functional specialization in segregation of neurons expressing distinct types of calcium-binding proteins is postulated.     

Primary retinal targets in the Atlantic loggerhead sea turtle,Caretta caretta

Summary Autoradiographic analysis distinguished twelve primary retinal targets in the diencephalon and the mesencephalon of the Atlantic loggerhead sea turtle, Caretta caretta. While the majority of fibers terminate contralaterally, sparse labelling is seen over ipsilateral thalamic nuclei. The dorsal optic nucleus is the most expansive retinal target in the dorsal thalamus. Four nuclei ventral and one dorsal, to the dorsal optic nucleus, receive retinal input. Before terminating in the optic tectum, labelled fibers pass through the pretectum terminating in four nuclei. Within the superficial zone of the optic tectum, three terminal zones are recognized. A distinct accessory tegmental tract separates from the main optic tract terminating in the basal optic nucleus.While such a multiplicity of retinal targets occurs among other reptiles, birds and mammals, it is presently impossible to accurately recognize visual homologies among amniotic vertebrates.     

Level of metabolic activity (cytochrome oxidase) as an index of functional significance of tectofugal and thalamofugal channels of the reptilian visual system

Using histochemical determination of activity of the mitochondrial oxidative enzyme cytochrome oxidase (CO) the metabolic activity in structures of brain of turtles and lizards was shown to be higher in centers of the tectofugal channel (tectal SGC, pretectal Ptv, thalamic Rot, telencephalic visual area of Advr of) than in the centers of the thalamofugal channel (thalamic GLd, Cxdl and Path of telencephalon) of the visual system. Some interspecies differences in distribution of CO activity in the tectal, thalamic, and telencephalic visual centers were revealed between terrestrial and pond turtles and lizards. The obtained data confirm the idea of the dominating role of the tectofugal visual pathway in information processing and organization of the usual behavior of reptiles.     

Calcium-binding proteins and metabolic activity in thalamotelencephalic parts of the turtle visual system

Distribution of three calcium-binding proteins (CaBPr) calbindin (CB), calretinin (CR) and parvalbumin (PV) in parallel with metabolic activity (cytochrome oxidase, CO) was studied in telencephalic projection zones of the tecto- and thalamofugal visual pathways in experiments on the Horsfield's terrapin Testudo horsfieldi and the pond turtle Emys orbicularis. It was shown that the nucleus rotundus (Rot) and dorsal lateral geniculate nucleus (GLd) terminal fields in both zones (dorsolateral region of the anterior ventricular ridge, Advrdl and dorsolateral cortex, Cxdl, respectively) were CB-immunoreactive (-ir) in the both studied turtle species. The highest density of CB-ir terminals and the focus of rotundal projections in the Advrdl core coincided precisely. The GLd terminal field in Cxdl also was CR-ir. The PV contribution to innervation of both projectional zones was much lower, especially to innervation of Cxdl from GLd. In spite of similar CB-ir innervation, the projectional field of the tectofugal pathway of Advrdl had the much higher CO activity than of that of the thalamofugal pathway in Cxdl. The neurons immunoreactive to all three CaBPr types were distributed in Cxdl in different ratios in each of layers. In the visual Advrdl area the overwhelming majority were PV-ir neurons, whereas CB-ir neurons were absent. The conclusion is made that in spite of the CB- or CB/CR-immunoreactivity predominates over the PV-immunoreactivity in both thalamotelencephalic pathways of the visual system, the tectofugal (rotundo-Advrdl) pathway having the higher metabolic activity.     

Afferent connections of the optic tectum in channel catfish Ictalurus punctatus

Summary Horseradish peroxidase was injected unilaterally into the optic tectum of the channel catfish, Ictalurus punctatus. The sources of tectal afferents were thereby revealed by retrogradely labeled neurons in various brain centers. Retrogradely labeled cells were seen in both the ipsilateral and contralateral telencephalon. The superficial pretectal area was labeled on both sides of the brain. Ipsilateral projections were also observed coming from the entopeduncular nucleus. Both the anterior thalamic nucleus and the ventro-medial thalamic nucleus projected to the ipsilateral optic tectum. Cells in the ipsilateral nucleus of the posterior commissure were seen to project to the tectum. Labeled fibers were visualized in the lateral geniculate nucleus ipsilateral to the injected tectum, however, no labeled cell bodies were observed. Therefore, tectal cells project to the lateral geniculate nucleus, but this projection is not reciprocal. No labeled cells were found in the cerebellum. Labeled cells occurred in both the ipsilateral and contralateral medial reticular formation; they were also observed in the ipsilateral nucleus isthmi. A projection was seen coming from the dorsal funicular nucleus. Furthermore, labeled cells were shown in the inferior raphe nucleus.Abbreviations AP Area pretectalis - C Cerebellum - DPTN Dorsal posterior tegmental nucleus - H Habenula - IRF Inferior reticular formation - LI Inferior lobe - LGN Lateral geniculate nucleus - LR Lateral recess - MB Mammillary body - MRF Medial reticular formation - MZ Medial zone of the telencephalon - NC Nucleus corticalis - NDL-M Nucleus opticus dorsolateralis/pars medialis - NI Nucleus isthmi - NPC Nucleus of the posterior commissure - OPT Optic tectum - OT Optic tract - PC Posterior commissure - PN Pineal organ - PrOP Preoptic nucleus - PT Pretectum - TBt Tectobulbar tract - TEL Telencephalon - TL Torus longitudinalis - TS Torus semicircularis - VC Valvula cerebelli - VLTN Ventrolateral thalamic nucleus - VMTN Ventromedial thalamic nucleus     

Efferent neurons of the auditory center in the midbrain of the frog Rana ridibunda

Individual cells which produce projections from the torus semicircularis in the frog have been visualized after injection of horseradish peroxidase (HRP) to various thalamic and isthmal areas. Labeled toral cells were observed if HRP had been injected to the posterodorsal areas of the thalamus or to the isthmal areas where lateral lemniscus fibers and cells of the premature lateral lemniscal nucleus are situated. Medium and large size cells in the rostrolateral torus semicircularis were mainly labeled. Thalamic injections of the HRP produced more labeled cells in the lateral part of the magnocellular nucleus, whereas isthmal injections produced labeled cells mainly in the lateral part of the laminar nucleus. A few HRP containing cells were observed in the principal nucleus of the torus. Specificity of the neuronal organisation of the auditory pathway in amphibians is discussed.     

Neurochemical organization of reptilian thalamus. Comparative analysis of amniote optical centers

Study of neurochemical characteristics of the turtle thalamus was performed using antibodies against several biologically active compounds; monoamines (5-HT, TH), neuropeptides (SP, m-Enk, NPY), against ChAT and by histochemical detection of NADPH-d. Based on our own results and literature data obtained on other representatives of reptiles, birds, and mammals, a comparative analysis was carried out of neurochemical organization of two thalamic optical centers—relay nuclei of thalamofugal (GLd) and tectofugal (Rot/LP-Pulv) systems of amniotes. Features of similarities and differences of these centers in representatives of non-mammalian and mammalian amniotes are revealed. GLd has a great similarity of the studied neurochemical characteristics in all amniotes. It receives innervation from all studied transmitter-modulatory systems with predominance of serotonin-, choline- and NPY-ergic projections. Neurochemical organization of Rot comparable with the tectorecipient part of mammalian LP-Pulv has a great resemblance in reptiles and birds, with considerable interspecies differences inside each class. In the majority of the studied systems, Rot is characterized by scant innervation. On the contrary, LP-Pulv receives sufficiently massive innervation from these systems. The most characteristic of this complex are rich SP- and cholinergic projections that are scant in Rot of reptiles and birds. The similar feature of Rot and LP-Pulv is the presence of massive serotonin- and NO-ergic (NADPH-positive) projections. The revealed similarities and differences of the neurochemical characteristics of thalamic optical centers among amniotes seem to reflect various transformations of thalamo- and tectofugal visual systems in the course of phylogenetic and adaptive evolution.     

Demonstration of afferent connections of the forebrain in Emys orbicularis turtles by the peroxidase method

The transport of horseradish peroxidase (HRP) out of the injection site in the dorsal ventricular ridge was studied in turtles Emys orbicularis. Labeled cells in the forebrain were observed in the paleostriatum among fibers of the lateral forebrain bundle. In the thalamus most of cells containing the granular HRP reaction product were located in the n. rotundus, n. reuniens and perirotundal nuclei (n. dorso-medialis anterior, n. magnocellularis thalami, n (centralis) lateralis, n. dorso-medialis). Fewer labeled cells were revealed in the n. anterior and n. ventralis. The density of labeled cells in the majority of all thalamic nuclei increased if the HRP was extended from the dorsal ventricular ridge into the neostriatum and the pallial thickening with adjacent general cortex. HRP positive cells in the pretectal area, nuclei of the posterior commissura and mesencephalic ventro-lateral tegmentum were observed only in cases when the enzyme was diffused from the injection site into the neostriatum, while the HRP retrograde transport to n. geniculatus lateralis, pars dorsalis was revealed only when HRP was extended into the pallial thickening and adjacent general cortex. Ascending connections of the paleostriatum, thalamic nuclei and mesencephalic tegmentum with telencephalic structures, mainly with the dorsal ventricular ridge, were discussed.     

Retinal projections in European Salamandridae

Summary The retinal projections to the brain were studied in three species of European Salamandridae using anterograde transport of horseradish peroxidase and autoradiography. The results obtained were basically identical for all species and confirmed earlier findings on the fiber supply to the preoptic nucleus and the basal optic neuropil. In the anterior thalamus projections to three distinct terminal fields are clearly visible: (i) the diffusely stained corpus geniculatum thalamicum, (ii) the neuropil of Bellonci, pars lateralis, and (iii) a dorsomedial terminal field, the neuropil of Bellonci, pars medialis. Caudal to these terminal fields is an almost terminal-free region, the lateral neuropil. In the posterior thalamus a medial terminal field, the uncinate field, and a laterally located terminal field, the posterior thalamic neuropil, are distinguishable. The tectum opticum displays as many as four dense layers of retinofugal fibers and terminals in the rostral part and, in addition, a more densely stained strip of neuropil running from rostral to caudal over the tectum. The extent of ipsilateral fibers is greater than previously reported in other urodele species. They supply the medial and the lateral parts of the neuropil of Bellonci, the uncinate field, and reach the tectum opticum via the medial optic tract. Further, they form terminals in the innermost optic fiber layer throughout the rostral half of the ipsilateral tectum. A small proportion of ipsilateral fibers contributes very sparsely to all other thalamic terminal fields, leaving only the caudal part of the tectum and several layers of the rostral tectum completely free of a direct retinofugal fiber supply.     

Primary projections of the trigeminal nerve in two species of sturgeon: Acipenser oxyrhynchus and Scaphirhynchus platorynchus

Horseradish peroxidase histochemical studies of afferent and efferent projections of the trigeminal nerve in two species of chondrostean fishes revealed medial, descending and ascending projections. Entering fibers of the trigeminal sensory root project medially to terminate in the medial trigeminal nucleus, located along the medial wall of the rostral medulla. Other entering sensory fibers turn caudally within the medulla, forming the trigeminal spinal tract, and terminate within the descending trigeminal nucleus. The descending trigeminal nucleus consists of dorsal (DTNd) and ventral (DTNv) components. Fibers of the trigeminal spinal tract descend through the lateral alar medulla and into the dorsolateral cervical spinal cord. Fibers exit the spinal tract throughout its length, projecting to the ventral descending trigeminal nucleus (DTNv) in the medulla and to the funicular nucleus at the obex. Retrograde transport of HRP through sensory root fibers also revealed an ascending bundle of fibers that constitutes the neurites of the mesencephalic trigeminal nucleus, cell bodies of which are located in the rostral optic tectum. Retrograde transport of HRP through motor root fibers labeled ipsilateral cells of the trigeminal motor nucleus, located in the rostral branchiomeric motor column.     

Retinofugal and retinopetal projections in the teleost Channa micropeltes (Channiformes)

Summary Retinofugal and retinopetal projections were investigated in the teleost fish Channa micropeltes (Channiformes) by means of the cobaltous lysine and horseradish peroxidase (HRP) tracing techniques. Retinofugal fibers cross completely in the optic chiasma. A conspicious lamination is present in those parts of the optic tract that give rise to the marginal branches of the optic tract. This layering of optic fibers continues in the marginal branches to mesencephalic levels. Retinal projections to the preoptic and hypothalamic regions are sparse; they are more pronounced in the area of pretectal nuclei. The medial pretectal complex and the cortical pretectal nucleus are more fully differentiated than in other teleostean species. Further targets include the thalamus and the optic tectum. The course of major optic sub-tracts and smaller fascicles is described. Retinopetal neurons are located contralaterally in a rostral and a caudal part of the nucleus olfactoretinalis, and in a circumscribed nucleus thalamoretinalis. The present findings are compared with reports on other teleost species.     

Retinofugal and retinopetal connections in the upside-down catfish (Synodontis nigriventris)

Summary The retinofugal and retinopetal connections in the upside-down catfish Synodontis nigriventris were studied by use of the horseradish-peroxidase (HRP) techniques, autoradiography, and degeneration-silver methods. An unusual retinal projection to the torus semicircularis as well as projections to the retina from three different sources in the brain are described. After intra-ocular injections of HRP, labeled cells were found in the optic tectum, the dorsomedial optic nucleus and one of the pretectal nuclei. These new findings support the basic hypothesis (i) that neuronal connections are more extensive in primitive brains, and (ii) that the evolutionary development of more complex brains involves the loss of some selected connections.     

DC electrical stimulation of the pretectal thalamus and its effects on the feeding behavior of the toad (Bufo bufo)

The feeding motivation of the common European common toad (Bufo bufo) can be quantified by the feeding sequence of arousal-orientation-approach-fixate-snap. Previous work has found that the optic tectum is an important structure responsible for the mediation of feeding behaviors, and combined electrical and visual stimulation of the optic tectum was found to increase the animals feeding behaviors. However, the pretectal thalamus has an inhibitory influence upon the optic tectum and its lesion results in disinhibited feeding behaviors. This suggests that feeding behavior of anurans is also subject to influence from the pretectal thalamus. Previous studies involving the application of DC stimulation to brain tissue has generated slow potential shifts and these shifts have been implicated in the modulation of the neural mechanisms associated with behavior. The current study investigated the application of DC stimulation to the diencephalon surface dorsal to the lateral posterodorsal pretectal thalamic nucleus in Bufo bufo, in order to assess effects on feeding motivation. The application of DC stimulation increased the incidence of avoidance behaviors to a visual prey stimulus while reducing the prey catching behavior component of approach, suggesting that the DC current applied to the pretectum increased the inhibition upon the feeding elements of the optic tectum. This can be explained by the generation of slow potential shifts.     

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.