Optic tectum of the eastern garter snake, Thamnophis sirtalis. III. Morphology of intrinsic neurons

Extracellular iontophoretic injections of horseradish peroxidase and Golgi preparations were used to study the distribution and morphology of intrinsic neurons of the garter snake optic tectum. Four morphologically distinct classes of neurons were identified. The type A neuron is found throughout the retinorecipient tectal layers. It has a large, fusiform soma and infrequently branching dendrites that radiate in the horizontal plane and are studded with varicose appendages. An axon arises from the soma or proximal dendrite and gives rise to widely spreading branches that overlap the cell's dendritic field. The type B neuron has a small, spherical soma in sublayer b of the stratum fibrosum et griseum superficiale. Thick, varicose dendrites ascend from the soma and form a bushy arbor in the overlying sublayer a. A thin axon descends vertically from the soma and arborizes in vertical alignment with the cell's dendritic field in sublayer c of the stratum fibrosum et griseum superficiale and the upper third of the stratum griseum centrale. The type C neuron is a bipolar cell with a small, vertically fusiform soma situated at the upper border of the stratum griseum centrale. Thin, sparsely branching dendrites extend vertically into the superficial and central gray layers. An axon arises from the soma and courses ventrally into the stratum griseum centrale where it gives rise to a plexus of widely spreading branches that extend medially from the cell's dendritic field. The type D neuron is a small, stellate cell with a spherical soma and fine, appendage-laden dendrites that are restricted to the stratum griseum centrale. The axon of the type D cell courses in the central gray where it gives rise to widely spreading branches that extend laterally from the cell's dendritic field.     

The optic tectum of the carp: pyramidal neurons of the SFGS

We have studied the stratum fibrosum et griseum superficiale (SFGS) of the carp optic tectum with optic and electron microscopy. This stratum is a dense neuropil with disordered appearance, in which numerous neuronal bodies of different characteristics and variable distribution according to the tectal regions are intercalated, more abundant in the dorsomedial zones of the tectum. Within these neuronal types, the most characteristic of SFGS are the large pyramidals of vertical development. Such neurons shows an ascendant dendritic shaft, very developed in the stratum marginale, a thinner dendritic shaft in the basal pole and a descending axon that reaches the internal zones of the stratum griseum centrale. Glial elements are highly associated to the pyramidal neuron bodies. The synaptic contacts are abundant and of various types, specially on the spinous dendritic branches which lie in the stratum marginale.     

Tectothalamic visual projections in turtles: their cells of origin revealed by tracing methods

In two species of turtle (Emys orbicularis and Testudo horsfieldi), retrograde and anterograde tracer techniques were used to study projections from the optic tectum to the nucleus rotundus (Rot) and to the dorsal lateral geniculate nucleus (GLd). The ipsilateral Rot received the most massive tectal projections, stemming from numerous neurons located in the stratum griseum centrale (SGC). These neurons varied in size and shape, many of them having a wide zone of dendritic arborization within both the (SGC) and the stratum griseum et fibrosum superficiale (SGFS). Projections from the tectum to the GLd were ipsilateral, were extremely scarce, and arose from a small number of neurons of various shapes situated in the SGFS; these cells were, as a rule, smaller than those projecting to the Rot. For the most part, these neurons were radially oriented, with rather restricted dendritic arborizations in the most superficial sublayers of the SGFS; smaller numbers of projection neurons were horizontally oriented, with long dendrites branching throughout the layer. Some neurons located in the stratum griseum periventriculare (SGP) projected to both the Rot and the GLd. Most of these neurons had dendritic arborizations within the retinorecipient zone of the SGFS. We were unable to rule out the possibility that some cells projecting to the GLd were situated in the SGC. Both the GLd and the main body of the Rot did not contain neurons projecting to the optic tectum. Thalamic neurons projecting to the tectum were observed in the ventral lateral geniculate nucleus, the intergeniculate leaflet and the interstitial nuclei of the tectothalamic tract, and the nucleus of the decussatio supraoptica ventralis. The question of whether variation in the laminar organization of the tectorotundal and tectogeniculate projection neurons in reptiles, birds, and mammals may be related to different degrees of differentiation of the tectal layers is discussed.     

Periventricular efferent neurons in the optic tectum of rainbow trout

The efferent connections and axonal and dendritic morphologies of periventricular neurons were examined in the optic tectum of rainbow trout to classify periventricular efferent neurons in salmonids. Among the target nuclei of tectal efferents, tracer injections to the following four structures labeled periventricular neurons: the area pretectalis pars dorsalis (APd), nucleus pretectalis superficialis pars magnocellularis (PSm), nucleus ventrolateralis of torus semicircularis (TS), and nucleus isthmi (NI). Two types of periventricular neurons were labeled by injections to the APd. One of them had an apical dendrite ramifying at the stratum fibrosum et griseum superficiale (SFGS), with an axon that bifurcated into two branches at the stratum griseum centrale (SGC), and the other had an apical dendrite ramifying at the SGC. Two types of periventricular neurons were labeled after injections to the TS. One of them had an apical dendrite ramifying at the boundary between the stratum opticum (SO) and the SFGS, and the other had dendritic branches restricted to the stratum album centrale or stratum periventriculare. Injections to the PSm and NI labeled periventricular neurons of the same type with an apical dendrite ramifying at the SO and a characteristic axon that split into superficial and deep branches projecting to the PSm and NI, respectively. This cell type also possessed axonal branches that terminated within the tectum. These results indicate that periventricular efferent neurons can be classified into at least five types that possess type-specific axonal and dendritic morphologies. We also describe other tectal neurons labeled by the present injections.     

Fiber connections of the torus longitudinalis and optic tectum in holocentrid teleosts

Fiber connections of the torus longitudinalis (TL) and target(s) of toral recipient tectal neurons (pyramidal cells) in the optic tectum were examined by tract-tracing methods in holocentrids. Injections into the stratum marginale (SM) labeled neurons in the stratum opticum and stratum fibrosum et griseum superficiale (SFGS). They had superficial spiny dendrites, with a fan-shaped branching pattern in SM and a thick basal dendrite that gave rise to bushy horizontal branches at the boundary between the SFGS and the stratum griseum centrale (SGC), where an axon and a thin dendrite arose. The axon terminated in a middle cellular layer of the SGC, and the thin dendrite ramified slightly deeper to this cellular layer. The SM injections also labeled cells in the ipsilateral TL. Injections into either the lateral or the medial part of TL labeled terminals in the ipsilateral SM and neurons in the bilateral nucleus paracommissuralis (NPC) and nucleus subvalvularis and ipsilateral nucleus subeminentialis. Only medial TL injections labeled cells in the ipsilateral SGC. These neurons had a basal dendrite that branched in the middle cellular layer of SGC, suggesting that they receive inputs from the pyramidal cells and project back to the TL to form a closed circuit. Only lateral TL injections labeled terminals in the corpus cerebelli. A visual telencephalic portion projects to the NPC and sublayers of SGC, where dendrites of the pyramidal cells and SGC neurons ramify. The present results therefore suggest that the TL and SM are components of an intricate circuitry that exerts telencephalic descending visual influence on the optic tectum and corpus cerebelli.     

Tectal projection neurons to the retinopetal nucleus in the filefish

Following horseradish peroxidase (HRP) injections into the preoptic retinopetal nucleus (PRN), neurons in the ipsilateral optic tectum were labeled retrogradely. Labeled neurons exhibited a 'Golgi-like' appearance, somata of these neurons were pyriform or round, and most of them were located in the stratum album centrale (SAC) or the stratum periventriculare (SPV). These neurons had a long apical dendrite, which ramified in the upper-half of SGC into horizontally arborized dendritic fields. The main trunk of the apical dendrites also gave off several branches in the stratum fibrosum et griseum superficiale (SFGS) and reached the stratum opticum (SO). These neurons resemble the 'large pyriform neurons' of Vanegas et al. (Vanegas, H., Laufer, M. and Amat, J., The optic tectum of a perciform teleost. I. General configuration and cytoarchitecture, J. Comp. Neurol., 154 (1974) 43-60) except that in the tecto-PRN neurons the axons originates from the apical dendritic shaft at or near the level of the SAC. Judging from their dendritic patterns, the tectal neurons projecting indirectly to the retina may receive non-retinal inputs besides the retinal input.     

Immunohistochemical localization of nicotinic acetylcholine receptor subunits in the mesencephalon and diencephalon of the chick (Gallus gallus).

Monoclonal antibodies against two alpha-bungarotoxin-binding subunits (alpha 7 and alpha 8) of the nicotinic acetylcholine receptors (nAChRs) were used as immunohistochemical probes to map their distribution in the chick diencephalon and mesencephalon. The distribution of the alpha 7 and alpha 8 nAChR subunits was compared to the distribution of immunoreactivity produced by a monoclonal antibody against the beta 2 structural subunit of the nAChRs. Structures that contained high numbers of alpha 7-like immunoreactive (LI) somata included the intergeniculate leaflet, nucleus intercalatus thalami, nucleus ovoidalis, organum paraventricularis, nucleus rotundus, isthmic nuclei, nucleus trochlearis, oculomotor complex, nucleus interstitio-pretecto-subpretectalis, stratum griseum centrale of the optic tectum, and nucleus semilunaris. Neuropil staining for alpha 7-LI was intense in the nucleus dorsomedialis hypothalami, nucleus geniculatus lateralis ventralis, griseum tecti, isthmic nuclei, nucleus lentiformis mesencephali, nucleus of the basal optic root, and stratum griseum et fibrosum superficiale of the tectum. High numbers of alpha 8-LI somata were found in the stratum griseum et fibrosum superficiale of the tectum and the nucleus interstitio-pretecto-subpretectalis, and intense neuropil staining for alpha 8-LI was found in the dorsal thalamus, nucleus geniculatus lateralis ventralis, lateral hypothalamus, griseum et fibrosum superficiale of the tectum. High numbers of beta 2-LI somata were found only in the nucleus spiriformis lateralis, whereas neuropil staining for beta 2-LI was intense in the nucleus geniculatus lateralis ventralis, nucleus suprachiasmaticus, nucleus lateralis anterior, nucleus habenularis lateralis, area pretectalis, griseum tecti, nucleus lentiformis mesencephalis, nucleus externus, and nucleus interpeduncularis, and in the stratum griseum centrale, stratum griseum et fibrosum superficiale, and stratum opticum of the tectum. These results indicate that there are major disparities in the localization of the alpha-bungarotoxin-binding alpha 7 and alpha 8 nAChR subunits and the beta 2 structural nAChR subunit in the chick diencephalon and mesencephalon. These nAChR subunits appear, however, to coexist in several regions of the chick brain.     

Tectoreticular pathways in the turtle, Pseudemys scripta. II. Morphology of tectoreticular cells

The morphology of tectoreticular neurons in turtles was examined with serial section reconstructions of neurons retrogradely filled with HRP. Six classes of tectal neurons project into the three tectobulbar pathways characterized in the preceding paper (Sereno, '85). (1) Large multipolar neurons with somata in the central gray layers, and with moderately branched dendrites sometimes spanning over a millimeter, project into the dorsal tectobulbar pathway, TBd. Their dendrites are covered with fine spicules and tend to arborize in the lower third of the superficial gray layers. (2) Medium-sized neurons with multiple radial dendrites and somata in the central white and upper periventricular layers probably project into the ipsilateral intermediate tectobulbar pathway, TBi. Their dendrites also bear fine spicules and usually reach the tectal surface. (3) Small radial cells in the periventricular layers, and (4) small bitufted radial cells in the superficial gray project into the small caliber component of the ipsilateral ventral tectobulbar pathway, TBv(sm). (5) Medium-sized central gray neurons with stratified dendrites, and (6) medium-sized central gray neurons with horizontal dendrites probably project into the medium caliber component of the ventral tectobulbar pathway, TBv(med). In contrast to TBd and TBi neurons, these last four classes emit a spray of long, filamentous dendritic appendages in the central gray and have dendritic arbors near the top of the superficial gray. The morphology of the neurons described in this and the preceding paper is briefly discussed in light of current ideas about tectally mediated sensorimotor transformations.     

Calretinin-like immunoreactivity in the optic tectum of the tench (Tinca tinca L.)

The distribution of calretinin-like immunopositive cells and fibers in the optic tectum of the tench (Tinca tinca) was studied by using a polyclonal antibody and the avidin-biotin-peroxidase technique. A clear laminated pattern of calretinin-like immunoreactivity was observed. The stratum periventriculare demonstrated a large number of strongly labeled cells whereas in the strata album centrale and griseum centrale, and at the boundary between the strata griseum centrale and fibrosum et griseum superficiale, some scarce, weakly immunostained cells were observed. No immunoreactive cells were seen in the strata fibrosum et griseum superficiale, opticum and marginale. Cells belonging to neuronal types X and XIV, previously characterized using Golgi impregnation, were found to be calretinin-like immunoreactive. Most calretinin-like immunopositive fibers were found in the strata fibrosum et griseum superficiale and opticum with a distribution pattern similar to retinotectal axons in these layers. In agreement with previous biochemical studies, our data suggest that, by contrast to all other classes of vertebrates, instead of calretinin and calbindin D-28k, only one protein is present in teleosts. Nevertheless, the calretinin-like immunostaining pattern in the teleost optic tectum was more complex than that previously described for calbindin D-28k. When compared to the calretinin-immunostaining in the rat superior colliculus, it is evident the presence in both amniotes and anamniotes of calretinin-immunopositive retinotectal axons. However, the distribution patterns of intrinsic calretinin-immunoreactive cells were different. Immunolabeled cells have been described in all layers of the superior colliculus, whereas the cells containing calretinin were restricted to the three deep strata of the tench optic tectum, a more similar distribution to what has been reported in the chick optic tectum.     

Quantitative histological studies of the optic tectum in six species of Notropis and Cyprinella (Cyprinidae, Teleostei).

Significant differences in stratification and size of the visual layers of the optic tectum were found between three clear-water minnows (Notropis amabilis, N. boops, Cyprinella venustas) and three turbid-water minnows (N. atherinoides, N. bairdi, and C. lutrensis). Correlations among a variety of neural structures suggested the importance of stratum marginale (SM), stratum opticum (SO), and stratum fibrosum et griseum superficiale (SFGS), stratum griseum centrale (SGC) and stratum periventriculare (SPV) in vision, of stratum album centrale (SAC) and SGC for olfaction, and of SPV for the processing of acoustico-lateral information.     

Glutamate-immunoreactivity in the retina and optic tectum of goldfish

Glutamate was immunohistochemically localized in the goldfish retina and tectum at the light and electron microscopic (E.M.) levels using double affinity purified antisera against glutaraldehyde conjugated L-glutamate. In retina, glutamate-immunoreactivity (Glu+) was observed in cone inner segments, cone pedicles, bipolar cells, a small number of amacrine cells and the majority of cells in the ganglion cell layer. The latter were shown to be ganglion cells by simultaneous retrograde labeling. Centrally, Glu+ was observed in axons in the optic nerve and tract, and in stratum opticum and stratum fibrosum et griseum superficialis (SFGS) of the tectum. The Glu+ in the optic pathway disappeared four days after optic denervation and was restored by regeneration without affecting the Glu+ of intrinsic tectal neurons. In tectum, Glu+ was also observed in torus longitudinalis granule cells, toral terminals in stratum marginale, some pyramidal neurons in the SFGS, multipolar and fusiform neurons in stratum griseum centrale, large multipolar and pyriform projection neurons in stratum album centrale, and many periventricular neurons. Glu+ was also localized within unidentified puncta throughout the tectum and within radially oriented dendrites of periventricular neurons. At the E.M. level, a variety of Glu+ terminals were observed. Glu+ toral terminals formed axospinous synapses with dendritic spines of pyramidal neurons. Ultrastructurally identifiable Glu+ putative optic terminals formed synapses with either Glu+ or Glu- dendritic profiles, and with Glu- vesicle-containing profiles, presumed to be GABAergic. These findings are consistent with the hypothesis that a number of intrinsic and projection neurons in the goldfish retinotectal system, including most ganglion cells, may use glutamate as a neurotransmitter.     

Diameters and terminal patterns of retinofugal axons in their target areas: an HRP study in two teleosts (Sebastiscus and Navodon)

Studies in various vertebrate classes, particularly amphibians and mammals, have revealed that retinal ganglion cells with different functional properties project by means of axons of correspondingly different diameters onto specific target regions. Whether a similar pattern exists in teleosts is partly investigated in the present study. HRP was injected into the optic nerve of Sebastiscus and Navodon. The calibers of intraretinal HRP-labeled axons were classed as fine (ca. 0.8 micron), medium (ca. 1.3 micron), and coarse (ca. 2.5 microns). The calibers of HRP-labeled retinofugal axons were then determined in their target areas, and these can be summarized as follows: Optic hypothalamus: fine, medium. Lateral geniculate nucleus: fine. Dorsolateral thalamic nucleus: fine, medium. Area pretectalis: fine. Nucleus of the posterior commissure: fine, medium. Area ventralis lateralis, contralateral: fine, medium, coarse; ipsilateral: coarse. Optic tectum, stratum opticum: fine, medium; stratum fibrosum et griseum superficiale: fine, medium, coarse, segregated in sublayers; stratum album centrale: fine, medium, coarse. Therefore, fine fibers were found to reach all target areas except the ipsilateral area ventralis lateralis, and these were the only fibers found in the lateral geniculate nucleus, area pretectalis, and stratum griseum centrale of the optic tectum. Coarse fibers, on the other hand, were found only in the area ventralis lateralis and the optic tectum (stratum fibrosum et griseum superficiale and stratum album centrale). Terminal patterns of these fibers were also studied. Most fine fibers take tortuous courses giving off a few branches and terminate with many varicosities, and medium and coarse fibers give off several finer branches and terminate with bulbous swellings. The physiological significance of these findings is discussed. In addition, retrogradely labeled (retinopetal) cells were found in the olfactory bulb and the area ventralis pars ventralis of the telencephalon, as well as in the preoptic area and the dorsolateral thalamic nucleus.     

Distribution of Met-enkephalin, Leu-enkephalin, substance P, neuropeptide Y, FMRFamide, and serotonin immunoreactivities in the optic tectum of the Atlantic salmon (Salmo salar L.)

The distribution of the neuropeptides methionine- and leucine-enkephalins, substance P, FMRFamide, neuropeptide Y, and vasoactive intestinal peptide, as well as the biogenic amine serotonin was studied in the optic tectum of the Atlantic salmon by means of immunocytochemistry. Peroxidase-antiperoxidase and indirect immunofluorescence methods were used to compare the differential laminar distribution of each of these substances. Nine parts of the optic tectum were selected for analysis on frontal sections: median, dorsolateral, and ventrolateral areas at rostral, medial, and caudal levels. Methionine- and leucine-enkephalin immunoreactive fibers were found in discrete sublayers in the following strata: stratum opticum, stratum fibrosum et griseum superficiale, stratum griseum centrale, stratum, and album centrale. Most of the substance P-, serotonin-, and vasoactive intestinal peptide-immunoreactive fibers were found in the stratum album centrale, whereas the FMRFamide- and neuropeptide Y-immunoreactive fibers were more or less randomly distributed within most of the strata of the optic tectum. Neuropeptide Y-immunoreactive cell bodies were located in the stratum periventriculare. We suggest an extrinsic origin for most of the immunoreactive fibers observed in the optic tectum, except for the neuropeptide Y-immunoreactive fibers that probably originate in the periventricular neurons. Although retinal peptidergic input to the optic tectum has been proposed in other vertebrates, there is no evidence that any of the neuropeptidelike or serotonin immunoreactive fibers in the optic tectum of the salmon should be of retinal origin. Differences and similarities with the distribution of neuropeptides in the optic tectum in representatives of other vertebrate classes are discussed.     

Distribution, laminar location, and morphology of tectal neurons projecting to the isthmo-optic nucleus and the nucleus isthmi, pars parvocellularis in the pigeon (Columba livia) and chick (Gallus domesticus): a retrograde labelling study.

Retrograde transport of Phaseolus vulgaris leucoagglutinin (PHA-L), fluorogold, fast blue, rhodamine labelled microspheres, and horseradish peroxidase (HRP) was employed to study the distribution, laminar location within the optic tectum, and morphology of tectal cells projecting upon the isthmo-optic nucleus (ION) and the nucleus isthmi, pars parvocellularis (Ipc), in the pigeon and chick. Following injections into the ION, all retrograde markers labelled tecto-ION neurons and their dendrites in the ipsilateral tectum. The cells were located within a relatively narrow band at the border between layers 9 and 10 of the stratum griseum et fibrosum superficiale (SGFS). Retrogradely labelled neuronal somata were different in both dendritic branching and shape; however, tecto-ION neurons generally possessed non-spiny radially oriented and multi-branched dendrites. The apical processes extended into the retino-recipient layers (2-7) of the SGFS and basal dendrites extended into layers 12-14 of the SGFS. Positive neuronal somata were observed throughout the rostro-caudal extent of the optic tectum. The average distance between adjacent tecto-ION neurons varied from one region to another. Specifically, retrogradely labelled cells were more numerous in the caudal, lateral, and ventral tectum, and less numerous at rostro-dorsal levels. Approximately 12,000 tecto-ION neurons were labelled within the ipsilateral optic tectum following either PHA-L or fluorescent dye injections. While the regional distribution of tecto-Ipc neurons was not examined, the morphology indicated that the cells had a single radially oriented dendritic process. Therefore, the apical dendrites are more restricted than those of tecto-ION cells. Moreover, the dendrites were spiny and arborized within layers 3, 5, and 9 of the ipsilateral optic tectum. The axon of tecto-Ipc cells arise from the apical process as a shepherd's crook and descend into the deep layers of the optic tectum. These results indicate that 1) tecto-ION and tecto-Ipc neurons are possibly monosynaptically activated by retinal input, 2) tecto-ION neurons are heterogeneous in morphology, and 3) there is a differential distribution of the tecto-ION neurons throughout the rostro-caudal extent of the optic tectum, suggesting a greater representation of the caudo-ventral portion of the optic tectum within the ION. The discussion primarily concerns the organization of the retino-tecto-ION-retinal circuit in light of the distribution and morphology of tecto-ION neurons within the optic tectum.     

Sonic motor nucleus and its connections with octaval and lateral line nuclei of the medulla in a rockfish, Sebastiscus marmoratus

The sonic motor nucleus and its fiber connections were examined in a rockfish, Sebastiscus marmoratus by means of tracer methods using horseradish peroxidase (HRP), biocytin, and carbocyanine dye (DiI). Sebastiscus has a swimbladder and a pair of extrinsic sonic/drumming muscles. The sonic muscle is ipsilaterally innervated by the occipital nerve which is composed of two ventral roots arising from the sonic motor nucleus. The sonic motor neurons are distributed in the most ventral part of the ventral column from the caudal medulla to the rostral spinal cord, and form a ventrally located columnar nucleus. Each neuron in this nucleus possesses a long thick dendrite and several short dendrites. The long dendrite extends dorsolaterally and branches in the lateral funiculus, whereas the short dendrites branch around their cell bodies. After biocytin injections into the sonic motor nucleus, two groups of premotor neurons were retrogradely labeled bilaterally, one in the dorsomedial portion of the descending octaval nucleus (DO) and the other in the medial zone of the reticular formation (RF) in the medulla. The DO premotor neurons were multipolar with several dendrites branching near the cell bodies, and the RF premotor neurons were bipolar. One of the two dendrites of the RF premotor neurons extends laterally into the ventral portion of the DO, and the other dendrite extends into the ventromedial area in the medulla. In the ventromedial dendritic field of the RF premotor neurons, descending fibers arising from the optic tectum (TO) and torus semicircularis (TS) traverse in the tractus tectobulbaris and terminate bilaterally. After DiI insertion into the ventromedial dendritic field, retrogradely labeled neurons were found bilaterally in the TS and TO. The majority of tectal neurons were located in the stratum griseum centrale. These neurons had two short basal dendrites branching in the cell layer and a long apical dendrite extending to the stratum fibrosum et griseum superficiale and stratum opticum. The toral neurons were bipolar and were distributed throughout the TS. Furthermore, biocytin injections into the medial nucleus of the lateral line system revealed that the nucleus projects bilaterally to the RF premotor neurons. These results show that premotor neurons for the sonic motor nucleus are located in the dorsomedial portion of the DO and the medial zone of the RF in the medulla. It is suggested that the sonic motor nucleus receives auditory input via the DO premotor neurons and input from RF premotor neurons which receive lateral line input via the medial nucleus, vestibular input through the lateral dendrite extending into the ventral portion of the DO, and information from the TO and TS via the tractus tectobulbaris.     

Optic tectum of the eastern garter snake, Thamnophis sirtalis. IV. Morphology of afferents from the retina

The morphology of single retinal terminals in the optic tectum of the eastern garter snake was demonstrated by orthograde filling from extracellular injections of horseradish peroxidase (HRP) into the optic tract. HRP-filled terminals share a characteristic shape and structure. Their parent axons course caudally in the stratum opticum within fascicles of 200-300 fibers of varying diameters. Single axons exit a fascicle and course into either the stratum fibrosum et griseum superficiale, ventrally, or the stratum zonale, dorsally, where they bifurcate successively two or three times into preterminal branches. Each preterminal branch gives rise to many thin, terminal branchlets laden with boutons. The arbors are ellipsoidal with their long axes oriented mediolaterally and their short axes oriented rostrocaudally. Arbors vary in their overall size (from 45 to 150 micron), in the diameters of their parent axons (from less than 0.5 to 3.0 micron), and in the size of their terminal boutons (from 0.5 to 3.5 micron). Bouton size increased with increasing diameter of the parent axon. The great majority of arbors are confined to one of three retinorecipient sublayers in the superficial tectum. However, the full range of arbor sizes and axon diameters is present in each sublayer.     

The projection of the retina upon the optic tectum of the pigeon.

The projection of the retina onto the optic tectum of the pigeon has been investigated using silver impregnation methods for degenerating axons and terminals, autoradiographic tracing, and the Golgi methods. Degenerating optic afferents distribute to sublaminae a-d and f of the stratum griseum et fibrosum superficiale over the whole tectum, but two major fields are observed. One occupies the caudal and ventral tectum (in which region laminar cytoarchitecture is marked), and the other rostral and dorsal tectum (where demarcation of cell laminae is relatively poor). Degeneration in the latter field is coarse and clearly distributes in a distinctly laminated fashion within the stratum griseum et fibrosum superficiale. In contrast, degeneration in the caudo-ventral region is fine, and laminated distribution less clear. The evolution of the degeneration pattern over survival periods from 3 to 56 days suggests that these laminar distributions reflect the existence of several different types of optic terminal ramification present in all parts of the tecum. A selective laminar distribution of such optic afferent types correlates well with our own and other Golgi studies. The results of the autoradiography experiments are broadly compatible with these findings.     

Comparative neurology of the optic tectum in ray-finned fishes: patterns of lamination formed by retinotectal projections

Retinotectal projections were studied in 33 different species of Actinopterygii, the ray-finned fishes, with horseradish peroxidase and cobalt tracing techniques. The distribution of retinorecipient layers in the contralateral optic tectum was analyzed. In addition, the degree of differentiation of the stratum periventriculate, and the presence of ipsilateral retinotectal projections was examined. Retinofugal fibers are labeled in the stratum opticum (SO), stratum fibrosum et griseum superficiale (SFGS), stratum griseum centrale (SGC), stratum album centrale (SAC) and stratum periventriculare (SPV). Some species lack the projection to the SO, others lack the projection to the SGC, and a third group of fishes lack both projections. Five different patterns of retinorecipient tectal strata are distinguished. These patterns correlate with the species' taxonomic position. Evolutionary trends of tectal lamination and retinotectal innervation are described. The retinotectal projection patterns provide a useful indicator of phylogenetic relationships. Some of our data suggest different relationships between actinopterygian species than hitherto believed.     

The visual connections of the adult flatfish, Achirus lineatus.

Metamorphosis in the flatfish is characterized by the migration of one eye around the dorsal surface of the head to a position adjacent to the other eye on the new top side of the animal. The visual connections of the adult flatfish, Achirus lineatus, were examined. Either the migrating or non-migrating eye was removed and the animal allowed to survive for one to three weeks. Alternate sections of the brain were stained by a modification of the Fink-Heimer technique, or with cresyl violet. The diencephalic visual connections of the flatfish were similar to those of other teleosts with contralateral projections to the nuclei corticalis, dorsomedialis thalami, pretectalis, and the corpus geniculatum laterale. The distribution of the retinal efferents to the optic tectum is unique in the flatfish. In the medial one-third of the tectum, terminal degeneration was found in three bands in the stratum opticum (SO) and the stratum griseum et fibrosum superficiale (sgfs). In the middle part of the tectum, two bands of degeneration remained over the sgfs. The lateral part of the tectum has only a very small amount of degeneration distributed radomly in scattered clusters over the deep SO and superficial sgfs. The Nissl preparations also reflected the differences between the medial and lateral parts of the tectum. Distinct layer was lacking in the medial tectum with a conspicuously absent large cell layer in the stratum griseum centrale (sgc). In contrast, the lateral tectum had a typical tectal stratification. Most notable were the large neurons of the sgc.     

Structural and functional characteristics of commissural neurons in the superior colliculus of the hamster

Intracellular recording and horseradish peroxidase (HRP) injection techniques were employed to delineate the structural and functional properties of superior collicular (SC) neurons in the hamster that were antidromically activated by electrical stimulation of the contralateral tectum. A total of 39 such cells were completely characterized, injected, and recovered. In ten of these, the axonal filling allowed us to reconstruct at least a portion of the terminal arborization in the SC contralateral to the labelled cell. Two of the recovered neurons were located in the stratum griseum superficiale (SGS), three were in the stratum opticum (SO), ten were in the stratum griseum intermediale (SGI), 11 were in the stratum album intermedium (SAI), 11 were in the stratum griseum profundum (SGP) and two were located in the stratum album profundum (SAP). The recovered cells were highly varied in both their morphological and their physiological characteristics. Somal areas ranged between 74 microns2 and 364 microns2, and the sample of recovered neurons included horizontal cells, narrow field vertical cells, and a variety of other multipolar neurons. Over one-third (38.5%) of the recovered cells were unresponsive, 2.6% were exclusively visual, 33.3% responded only to innocuous cutaneous stimuli, 10.2% were bimodal, 7.7% were specifically nociceptive, and 7.7% had complex (Rhoades, Mooney, and Jacquin: J. Neurosci. 3:1342-1354, '83) somatosensory receptive fields. We observed no clear-cut correlations between the structural and functional characteristics of these neurons. The conduction latencies of the commissural SC neurons ranged between 0.8 and 14.0 ms. The most rapidly conducting cells were located in the SGP and SAP. Conduction latency had a significant negative correlation with soma area. Labelled axons, in many cases, had at least one terminal arbor in a portion of the SC that was mirror symmetric with the location of the cell from which it originated. In several cases, however, commissural axons gave off a number of collaterals across the mediolateral extent of the tectum. commissural axonal terminations were visible only in the laminae ventral to the SO. Several commissural SC neurons also had extensive ipsilateral axon collaterals. Both the ipsilateral and commissural axon branches of these cells gave off en passant and terminal swellings.