Tectobulbar Tract Research Articles

Gamma Oscillations Are Generated Locally in an Attention-Related Midbrain Network

Gamma-band (25-140 Hz) oscillations are a hallmark of sensory processing in the forebrain. The optic tectum (OT), a midbrain structure implicated in sensorimotor processing and attention, also exhibits gamma oscillations. However, the origin and mechanisms of these oscillations remain unknown. We discovered that in acute slices of the avian OT, persistent (>100 ms) epochs of large amplitude gamma oscillations can be evoked that closely resemble those recorded in vivo. We found that cholinergic, glutamatergic, and GABAergic mechanisms differentially regulate the structure of the oscillations at various timescales. These persistent oscillations originate in the multisensory layers of the OT and are broadcast to visual layers via the cholinergic nucleus Ipc, providing a potential mechanism for enhancing the processing of visual information within the OT. The finding that the midbrain contains an intrinsic gamma-generating circuit suggests that the OT could use its own oscillatory code to route signals to forebrain networks.

Genetic Single-Cell Mosaic Analysis Implicates ephrinB2 Reverse Signaling in Projections from the Posterior Tectum to the Hindbrain in Zebrafish

The optic tectum is a visual center in vertebrates. It receives topographically ordered visual inputs from the retina in the superficial layers and then sends motor outputs from the deeper layers to the premotor reticulospinal system in the hindbrain. Although the topographic patterns of the retinotectal projection are well known, it is not yet well understood how tectal efferents in the tectobulbar tract project to the hindbrain. The retinotectal and the tectobulbar projections were visualized in a zebrafish stable transgenic line Tg(brn3a-hsp70:GFP). Using a single-neuron labeling system in combination with the cre/loxP and Gal4/UAS systems, we showed that the tectal neurons that projected to rhombomeres 2 and 6 were distributed with distinctive patterns along the anterior-posterior axis. Furthermore, we found that ephrinB2a was critically involved in increasing the probability of neurons projecting to rhombomere 2 through a reverse signaling mechanism. These results may provide a neuroanatomical and molecular basis for the motor command map in the tectum.

Connections of eye‐saccade‐related areas within mesencephalic reticular formation with the optic tectum in goldfish

Physiological studies demonstrate that separate sites within the mesencephalic reticular formation (MRF) can evoke eye saccades with different preferred directions. Furthermore, anatomical research suggests that a tectoreticulotectal circuit organized in accordance with the tectal eye movement map is present. However, whether the reticulotectal projection shifts with the gaze map present in the MRF is unknown. We explored this question in goldfish, by injecting biotin dextran amine within MRF sites that evoked upward, downward, oblique, and horizontal eye saccades. Then, we analyzed the labeling in the optic tectum. The main findings can be summarized as follows. 1) The MRF and the optic tectum were connected by separate axons of the tectobulbar tract. 2) The MRF was reciprocally connected mainly with the ipsilateral tectal lobe, but also with the contralateral one. 3) The MRF received projections chiefly from neurons located within intermediate and deep tectal layers. In addition, the MRF projections terminated primarily within the intermediate tectal layer. 4) The distribution of labeled neurons in the tectum shifted with the different MRF sites in a manner consistent with the tectal motor map. The area containing these cells was targeted by a high-density reticulotectal projection. In addition to this high-density topographic projection, there was a low-density one spread throughout the tectum. 5) Occasionally, boutons were observed adjacent to tectal labeled neurons. We conclude that the organization of the reticulotectal circuit is consistent with the functional topography of the MRF and that the MRF participates in a tectoreticulotectal feedback circuit.

Differential Responsiveness to the Chemorepellent Semaphorin 3A Distinguishes Ipsi- and Contralaterally Projecting Axons in the Chick Midbrain

In the chick dorsal mesencephalon, the optic tectum, the developing axons must choose between remaining on the same side of the midline or growing across it. The ipsilaterally projecting axons, forming the tectobulbar tract, course circumferentially toward the ventrally situated floor plate but before reaching the basal mesencephalon, the tegmentum, gradually turn caudally. Here, they follow the course of the medial longitudinal fasciculus (MLF), located parallel to the floor plate. By in vivo labeling of tectal axons, we could demonstrate that these axons arise primarily in the dorsal tectum. To test the idea that chemorepellent molecules are involved in guidance of the nondecussating axons, we performed coculture experiments employing tectal explants from various positions along the dorso-ventral axis. Axons emanating from dorsal tectal explants were strongly repelled by diencephalic tissue containing the neurons that give rise to the MLF whereas ventral tectal axons showed only a moderate response. This inhibitory effect was substantially neutralized by the addition of anti-neuropilin-1 antibodies. A similar differential response of axons was observed when tectal explants were cocultured with cell aggregates secreting the chemorepellent Semaphorin 3A (Sema3A). Sema3B and Sema3C, respectively, did not inhibit growth of tectal axons. In addition, neither the floor plate nor Slit2-secreting cell aggregates influenced outgrowth of dorsal fibers. In Sema3A-deficient mice, DiI-labeling revealed that dorsal mesencephalic axons cross the MLF instead of turning posteriorly upon reaching the fiber tract, thus behaving like the ventrally originating contralaterally projecting axons. A differential responsiveness of tectal axons to Sema3A most likely released by the MLF thus contributes to pathfinding in the ventral mesencephalon.

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

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

Distribution and substrate properties of agrin, a heparan sulfate proteoglycan of developing axonal pathways

The distribution and substrate properties of agrin, an extracellular matrix heparan sulfate proteoglycan (HSPG), was investigated in the developing chick nervous system by immunocytochemistry, Western blotting, and in neurite outgrowth assays. By comparing the distribution of agrin with that of laminin-1, merosin (laminin-2), neurofilament, and neural cell adhesion molecule (NCAM), it was found that throughout development, agrin is a constituent of all basal laminae. From embryonic day (E) 4 onwards, agrin is also abundant in axonal pathways of the central nervous system, such as the optic nerve, the tectobulbar pathway, the white matter of the spinal cord, and the marginal and the molecular layers of the forebrain and the cerebellum. The abundance of agrin in brain decreases from E13 onwards. In the peripheral nervous system, agrin is present throughout development as a constituent of the Schwann cell basal laminae. Western blots confirmed the immunocytochemical data, showing maximum expression of agrin occurs during the early to medium stages of brain development. Western blots also showed that in mouse and human brain, agrin exists as an HSPG. Purified agrin did not support neurite outgrowth, rather it inhibited retinal neurite extension on mixed agrin/merosin substrates. Despite the fact that agrin, when used as a substrate inhibited neurite outgrowth, its temporal and spatial overlap with growing axons suggests that agrin has a supportive role in the development of axonal pathways, possibly as a binding component for growth factors and cell adhesion proteins.

A mesencephalic relay for visual inputs to reticulospinal neurones in lampreys

Visual stimuli elicit motor responses in lampreys. These responses rely, in part, on the activation of reticulospinal (RS) neurones which constitute the main descending pathway in these early vertebrates. This study sought to identify and characterize possible mesencephalic relays for visual inputs to RS neurones of the rhombencephalon. The anatomical substrate subserving this function was investigated by iontophoretically ejecting cobalt-lysine, a retrograde tracer, in the middle rhombencephalic reticular nucleus in the in vitro isolated brainstem preparation of young adult Petromyzon marinus. Several populations of cells were retrogradely labeled in the brainstem. Of particular interest were the cell populations found on each side of rostral mesencephalon, located in the tectum and pretectum. There were, on average, 113 cells labeled contralateral to the injection site and 80 cells labeled ipsilateral to the injection site. The cells were morphologically similar on both sides, except that the contralateral group had larger cell bodies as compared to the group on the ipsilateral side. To determine whether the axons of the cells contacted reticulospinal neurones, electrophysiological experiments were carried out in which the region containing these cells was microstimulated. Large post-synaptic potentials were recorded intracellularly in RS neurones. Furthermore, microstimulation of the optic nerve on the same side as the recorded cell (i ON) evoked responses with a pattern similar to those resulting from stimulation of the optic tectum contralateral to the cell recorded (co OT), except for the longer response latencies. Local ejection of xylocaine (1% lidocaine hydrochloride) or CNQX (1 mM) onto the co OT reversibly abolished the responses evoked from stimulation of the i ON. There were no significant effects observed when the drug was ejected onto optic tectum ipsilateral to the cell. Taken together, the results from this study indicate that the crossed tectoreticular pathway is involved in relaying optic nerve inputs to RS neurones of the middle rhombencephalic reticular nucleus. Moreover, cells of origin of this pathway appear, in all respect, homologous to cells giving rise to the crossed tectobulbar pathway in other vertebrates.

Laminin alpha 2 chain (M chain) is found within the pathway of avian and murine retinal projections

Laminin-1 is found at the end-feet of neuroepithelial cells along the outer margin of the optic pathway during early stages of development. Prior to the establishment of most retinal projections in vivo, laminin-1 expression becomes restricted to basement membranes associated with the eye and optic pathway. We report that, in contrast to the alpha 1, beta 1, and gamma 1 chains of laminin-1, laminin alpha 2 chain (formerly laminin M chain) is expressed within the pathway of avian and murine retinal ganglion cell (RGC) growth cones as they extend into the optic nerve, across the optic chiasm and into the brain. Expression of laminin alpha 2 chain is reduced soon after formation of the visual projections but nevertheless maintained at non-basal lamina sites within the adult optic nerve. Laminin alpha 2 chain, in contrast to laminin-1 chains, is also highly expressed in the developing avian tectobulbar pathway. Chick optic nerve derived type-1 astrocytes in culture express laminin alpha 2 chains as extracellular fibrils on their surface. Laminin alpha 2 chain was also detected on the surface of cultured embryonic retinal neurons and developing RGCs. These results suggest that astrocytes and/or RGCS may synthesize laminin alpha 2 chain along the developing optic pathway, and imply that laminin alpha2--in a complex with non-beta1 and non-gamma1 laminin chains-may serve as an adhesive substrate and possibly as a guidance cue for elongating RGC growth cones in vivo.

Nucleus isthmi in goldfish: in vitro recordings and fiber connections revealed by HRP injections.

Recordings of field potentials in nucleus isthmi (NI) were obtained in an in vitro preparation of goldfish brain using a lateral approach. Horseradish peroxidase (HRP) was injected from recording electrodes to verify recordings within the nucleus and to label axonal pathways and cell bodies. Activity in NI was repetitive and could be elicited by stimulation of the optic nerve, tectum, pretectum, or tectobulbar tract. Spontaneous activity was present in some preparations and consisted of bursts with intervening silent periods. Anatomical and electrophysiological evidence indicated that the primary isthmotectal pathway is composed of fine fibers that exit NI rostrally and pass through pretectum to enter tectum rostrally. An afferent pathway consisting of both fine- and large-diameter fibers entered NI ventromedially; the large diameter axons have been previously reported in percomorph fishes, but were not thought to be present in cyprinids such as goldfish. The large diameter axons arise from labeled cell bodies in the region of the lateral thalamic nucleus. No labeled cell bodies were seen in ipsilateral nucleus pretectalis superficialis, pars magnocellularis, where they are seen in percomorphs. The fine axons, which have not been reported in percomorph fishes, were shown to arise from tectal bipolar (type VI) neurons. As in percomorphs, tectal type XIV neurons were also labeled. This and corroborating recordings from nucleus isthmi constitute the fist demonstration of a tectoisthmic projection in a cyprinid fish.

The long latency component of retinotectal transmission: Enhancement by stimulation of nucleus isthmi or tectobulbar tract and block by nicotinic cholinergic antagonists

The optic tectum of teleosts contains high concentrations of nicotinic and muscarinic acetylcholine receptors and receives putative cholinergic innervation from both nucleus isthmi in the tegmentum and a population of intrinsic tectal cells. Using in vitro preparations of goldfish brain, we have examined the effects of cholinergic antagonists and stimulation of nucleus isthmi on the tectal response to optic nerve stimulation. Our results show that: (1) a long latency component of the retinotectal field potential is polysynaptic in origin and occurs in isolated tectum; (2) this component can spread across the tectum from a beam of stimulated fibers and can appear in areas where the monosynaptic response is small or absent; (3) both monosynaptic and long latency components of the field potential are enhanced by prior stimulation of nucleus isthmi or the tectobulbar tract (15–300 ms); (4) both the long latency component of the field potential and the effects of stimulation of nucleus isthmis or tectobulbar tract are blocked by low concentrations of nicotinic antagonists; and (5) in deeper tectum a second polysynaptic response uncovered by pharmacological block of inhibition is not blocked by nicotinic antagonists. These results indicate that the cholinergic neurons intrinsic to tectum have a role in the spread of retinotectal excitation by nicotinic actions, and that stimulation of nucleus isthmi or tectobulbar tract facilitates activity in this system. There is in addition a separate recurrent excitatory circuit in tectum.

The avian tectobulbar tract: development, explant culture, and effects of antibodies on the pattern of neurite outgrowth

The tectobulbar tract is the first long-distance projecting fiber pathway to appear during the development of the avian optic tectum (dorsal half of the mesencephalon). Immunologically stained wholemounts of the E3 mesencephalon reveal that the early tectobulbar axons course in a dorsal-to-ventral direction and abruptly turn in a caudal direction shortly before reaching the ventral midline. During subsequent development, more tectobulbar axons are generated that form a parallel array of thick fascicles coursing ventrally within the mesencephalon. At this later stage of development, the tectobulbar tract bifurcates into an ipsilateral and contralateral pathway, both growing in a caudal direction near the mesencephalic ventral midline. Bifurcation and change in direction of growth is accompanied by a complete loss of the fasciculated growth pattern. Each tectobulbar axon is thus divided into a proximal fasciculated and a distal unfasciculated segment. Tectobulbar fascicles occupy the most superficial surface layer of the mesencephalon at early stages and are displaced deeper into the tissue beginning at embryonic day 5. In both of these locations, tectobulbar axons express molecules involved in axon-axon and axon-substrate interactions like the G4 antigen, neural cell adhesion molecule (N-CAM), neurofascin, and T61 antigen as revealed by immunohistochemistry and immunoblotting. Stripes of the mesencephalon explanted onto a basal lamina substratum show vigorous outgrowth of neurites. These processes grow in fascicles at a growth rate of 40 microns/h. Staining of the neurites with specific antibodies, as well as the position of the retrogradely labeled cell bodies, is in agreement with these processes being tectobulbar axons. This in vitro explant system was used to investigate the expression and possible functional involvement of N-CAM, neurofascin, G4 protein, and T61 antigen in the growth of these axons. The presence of antigen-binding fragments of polyclonal anti-G4 antibodies completely blocks fasciculation of the neurites but has no influence on their rate of elongation. Antibodies against N-CAM and neurofascin have no detectable effects. The number and length of the in vitro growing axons are reduced by the monoclonal T61 antibody. This effect is reversible. The elucidation of the exact course in vivo and the accessibility to the axons growing in vitro make the tectobulbar tract an excellent model system for the investigation of the role of these and other proteins in axonal growth and guidance during the development of the CNS.

A light- and electron-microscopic investigation of the optic tectum of the frog, Rana pipiens, II: The neurons that give rise to the crossed tecto-bulbar pathway

The superficial layers of the frog's optic tectum, Potter's (1969) layers A-G, comprise a complex neuropil made up of many afferent axons, the somata of a few neurons, and many dendrites from the neurons located in the deeper layers. Different types of retinal axons are believed to terminate in different layers (Maturana et al., 1960; Kuljis & Karten, 1988; Sargent et al., 1989), but little is known about the relationships between each type of input and the dendrites of the deep tectal neurons that extend into these superficial layers. The present study used the method of retrograde transport of horseradish peroxidase to study the synaptic contacts on the dendrites of the neurons that give rise to the crossed tecto-bulbar pathway. These cells have apical dendrites that ascend through the superficial retino-recipient layers. The somata of the cells that give rise to the crossed tecto-bulbar pathway are located in the superficial half of layer 6, preferentially clustered along the caudal, lateral, and rostral margins of the tectum. The somata of these cells range from 8-30 microns in diameter. Their axons are large (2-4 microns in diameter) myelinated fibers that arise from either their somata or proximal dendrites. Their axons travel within the deep medullary layer to leave the tectum at the lateral margin. Their dendritic arbors extend obliquely through the superficial layers to reach layer B where they turn and extend within the layer for up to 0.5 mm. The somata of these cells receive only a scant synaptic input. In contrast, their dendrites receive input in every layer, but the nature of this input varies from layer to layer. Synaptic terminals that resemble retinal ganglion cell boutons contact the labeled dendrites in layers B, F, and G. This indicates that the dendrites may receive monosynaptic input from several types of retinal ganglion cells. Terminals with small, flattened vesicles also contact the dendrites of these cells in each layer. In layer F and below, the terminals with flattened vesicles constitute 15% of the contacts; above layer F they constitute only 5-8% of the contacts. Terminals with medium-sized, flattened vesicles also contact the dendrites of these cells in every layer and constitute a large proportion of their input (33-95%). The latter terminals resemble those that are often postsynaptic to retinal terminals.

The first appearance of the future cerebral hemispheres in the human embryo at stage 14.

Thirty-five embryos of stage 14 (32 days) were studied in detail and graphic reconstructions of four of them were prepared. Characteristic features of this stage include the beginning formation of the future cerebral hemispheres and the cerebellar plates. The ventral boundary between telencephalon medium and diencephalon is the preoptic recess. Although a velum transversum is not yet distinguishable as a dorsal boundary, its site is indicated by a change in the thickness of the roof of the forebrain. As the cerebral vesicles (future hemispheres) begin to evaginate, a di-telencephalic sulcus and a corresponding lateral ventricle and ventricular ridge (torus hemisphericus) develop. The telencephalic wall is mainly ventricular layer but three areas show advanced differentiation: olfactory area, future amygdaloid body (which lies at first mainly in the diencephalon), and primordium of the hippocampus. The telencephalon is growing in length, and the forebrain now occupies almost one quarter of the total length of the brain. The two neuromeres of the diencephalon are no longer as clearly delineated. The floor of D1 presents a thickened chiasmatic plate; that of D2 includes the infundibulum, which is closely related to the adenohypophysial pouch. The ventricular surface of D1 presents elevations for the dorsal and ventral thalami, separated by the sulcus medius. Other features of the diencephalon include the ventricular eminence (medial ventricular ridge) of the basal nuclei and the hypothalamic cell cord, from which the preopticohypothalamotegmental tract arises. The roof of D2 contains the evaginating part of the synencephalon. The mesencephalic angle continues to diminish. Two neuromeres, M1 and M2, are still distinguishable. The oculomotor nucleus emits nerve fibres, as does also the trochlear nucleus, which lies in the isthmic segment. Some extracerebral oculomotor fibres are present, but decussating and extracerebral trochlear fibres have not yet appeared. In the region of the tectum, two nuclei are discernible, and will form the medial tectobulbar tract and the mesencephalic root of the trigeminal nerve, respectively. The medial longitudinal fasciculus is present. A "median ventricular formation" is sometimes found in the mesencephalic roof. The cerebellum is the widest part of the brain. Two neuromeres (isthmic segment and Rh1) are involved in its formation. Most of the cerebellar plate has differentiated an intermediate layer, and the future rhombic lip is discernible. Indications of an efferent fibre system are present. In addition to the cerebellum, the rhombencephalon includes Rh1 to Rh7, and RhD.(ABSTRACT TRUNCATED AT 400 WORDS)

Tectal neurons of the frog: intracellular recording and labeling with cobalt electrodes.

With the aid of cobalt-filled intracellular electrodes, responses evoked by electrical and visual stimulation were recorded from tectal neurons. Diffuse light and "edge" and "spot" stimuli were used for visual stimulation; electrical stimuli were delivered by means of bipolar electrodes to the optic tract. The cells were subsequently filled with cobalt for a correlative histological study. The following types of neurons were distinguished: (1) Pear-shaped neurons with restricted and broad dendritic fields occurring in tectal layers 8, 6, 4, and 2. In one group, the dendrites bore beads and small protrusions and the short axon arborized within, or close to, the dendritic arbor. They responded mostly with inhibitory postsynaptic potentials to the various stimulations. This type of neuron was interpreted to be a tectal interneuron. In the second group the dendrites were smooth, the dendritic arbor was large, and the axon could be followed to layer 7 in a few cases. Excitatory activities with frequent spike discharges prevailed in their responses. They were regarded as efferent neurons of the tectum. (2) Large ganglionic neurons showed a greater variation in form than has been described in previous Golgi studies. Their responses to the different stimulations were also very variable. The axons of five neurons could be followed to the dorsal part of the thalamus, to the postoptic commissural system, and to the direct and crossed tectobulbar tract. These axons emitted several collaterals to tectal, pretectal, and tegmental structures. (3) A single stellate neuron was impaled in layer 9. It responded only to electrical stimulation with excitation. (4) Deep bipolar neurons were not mentioned in earlier works. Their somata were found in layer 6; the vertically oriented dendrites arborized in plexiform layers 7, 5, and 3, over a distance of more than 1 mm. These cells were sensitive only to electrical stimulation.

Optic tectum of the eastern garter snake, Thamnophis sirtalis. II. Morphology of efferent cells.

Tectal efferent neurons were retrogradely filled from extracellular injections of horseradish peroxidase (HRP) into pathways efferent from the tectum. Tectorotundal neurons have cylindrical dendritic trees, 80-100 microns in diameter, that extend vertically across the central and superficial tectal layers. Apical and basal dendrites are laden with complex appendages. The axon gives rise to an intratectal, collateral arbor that extends horizontally into the stratum griseum centrale beyond the cell's dendritic tree. The parent axon exits the tectum laterally in the tectothalamic tract. Tectogeniculate neurons also have narrow, radially oriented, and highly branched apical dendrites, but their basal dendrites are infrequently branched and lack appendages. An intratectal axon collateral forms a small, spherical arbor overlapping the apical dendrites in sublayer c of the stratum fibrosum et griseum superficiale. The parent axon ascends vertically and just below the stratum opticum turns rostrad to follow the optic fibers to the diencephalon. Tectoisthmi neurons have small somata and thin, radial dendrites that arborize below the pial surface in the stratum zonale. An intratectal axon collateral forms a spatially restricted arbor ventral to the soma in register with the dendritic tree. Tectoisthmobulbar neurons have dendrites that arborize extensively in sublayer a of the stratum fibrosum et griseum superficiale. The axon exits the tectum without collateralizing and joins a small-caliber component of the ventral tectobulbar tract. Ipsilateral tectobulbar neurons have stellate dendritic fields, 150-250 microns in diameter, that are restricted to the deep layers of the tectum. Sparsely branched dendrites are appendage-free but bear many short, fine spicules. The axon initially ascends from the soma and recurves into the stratum album centrale without collateralizing before joining a medium-caliber component of the ventral tectobulbar tract. Crossed tectobulbar neurons have large, stellate dendritic trees with diameters ranging from 200 to 500 microns. Like ipsilateral tectobulbar neurons, their dendrites are appendage-free but bear spicules. Their thick-caliber axons exit the tectum without collateralizing and course deep in the stratum album centrale to reach the dorsal tectobulbar tract.

Optic tectum of the eastern garter snake, Thamnophis sirtalis. I. Efferent pathways.

Extracellular, iontophoretic injections of horseradish peroxidase were used to anterogradely fill axons efferent from the optic tectum in garter snakes. The tectal efferent pathways consist of six axon types with distinct projections and terminal morphologies. Tectogeniculate axons pass into the diencephalon via the optic tract, bearing collaterals that form spatially restricted, rodlike arbors in the pretectum, the ventral lateral geniculate nucleus, and the ventrolateral nucleus. Tectoisthmi axons exit the tectum as a thin-caliber component of the ventral tectobulbar tract. They form spatially restricted, spherical arbors within nucleus isthmi. Tectoisthmobulbar axons also give rise to small, spherical arbors within nucleus isthmi, but the parent axons continue caudally into the pontine and medullary reticular formation issuing many short collateral branches. Tectorotundal axons reach the diencephalon via the tectothalamic tract and give rise to fine terminal collaterals in the nucleus of the tectothalamic tract ipsilaterally and in nucleus rotundus bilaterally. Single axons form sheetlike terminal fields that span the rostrocaudal extent of nucleus rotundus. Ipsilateral tectobulbar axons descend into the midbrain tegmentum where they issue several thick collaterals that terminate widely throughout the nucleus lateralis profundus mesencephali. The parent axon continues caudally giving off several widely spreading collaterals within the pontine and medullary reticular formation. Crossed tectobulbar axons enter the dorsal tectobulbar tract and cross the midline to form the predorsal bundle. Single axons give rise to terminal collaterals in the nucleus lateralis profundus mesencephali bilaterally, the contralateral pontine and medullary reticular formation, and the intermediate gray of the cervical spinal cord.

Identification of a nucleus isthmi in the weakly electric fish Apteronotus leptorhynchus (Gymnotiformes).

The nucleus isthmi of teleost fish, amphibians, reptiles and birds, and its probable homologue, the nucleus parabigeminalis of mammals, share in common certain features such as location in the dorsal tegmentum and reciprocal connectivity with the optic tectum. In gymnotid fish the nucleus isthmi is located dorsolaterally in the brainstem tegmentum, ventral to the torus semicircularis and the lateral mesencephalic reticular area and dorsal to the rostral nucleus praeeminentialis. The nucleus isthmi has an ovoid shape, with a compact cellular part on its dorsal, medial and ventral aspects surrounding a hilar region with a sparse population of larger cells. Following wheat germ agglutinin-conjugated horseradish peroxidase injections into the optic tectum, anterogradely labeled fine terminals were observed leaving the tectobulbar tract and entering the ipsilateral nucleus isthmi via its laterally facing hilar region. Retrogradely labeled cells were present in the nucleus isthmi on both sides, indicating the presence of a bilateral isthmotectal projection similar to that reported in amphibians. The putative isthmal nucleus stains densely for acetylcholinesterase. Based on the similarity of its location, shape, cholinesterase histochemistry and reciprocal connectivity with the optic tectum, we identified this structure as the nucleus isthmi of gymnotids. An interesting observation of this study was that the nucleus isthmi, in addition to receiving fine terminals from the optic tectum, is also the recipient of a sparser population of thicker-caliber afferent fibers which terminate not only in the large-celled hilar region but also within the smaller-celled component of the nucleus; this projection appears to emanate from the torus semicircularis dorsalis.

A note on a tectal neuron projecting via the tectobulbar tract in teleosts

Catfish and goldfish received small, unilateral tectal injections of horseradish peroxidase. In these fish a population of large diameter tectal efferent axons could be seen travelling to the hindbrain. Each of these axons gave up to 19 collateral branches. These branches arborized in or near the dendritic fields of the ipsilateral nucleus lobobulbaris and oculomotor nucleus, and the contralateral trigeminal nucleus, oculomotor nucleus, abducens nucleus, nucleus lateralis valvulae and medial reticular formation. The results suggest that these axons are involved in triggering eye movements.

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.

Pathway choice by regenerating optic fibers following tectal lobectomy in the goldfish: Inferences from the study of gliosis in tectal efferent bundles

Cell counts in various fiber bundles of the goldfish brain have demonstrated a profound, but transient gliosis of tectal (and pretectal) efferent pathways following removal of a tectal lobe. In the majority of cases, the pathways which underwent gliosis were also those which were penetrated by regenerating optic fibers which had been sectioned by the tectal surgery. However, the dorsal trunk of the horizontal commissure on the intact side of the brain showed only a minimal gliotic response but was consistently innervated by the optic fibers. Conversely, the ansate commissure and the crossed tectobulbar tract invariably demonstrated a marked gliotic response but only rarely received more than minimal innervation by the regenerating fibers. These observations are discussed with regard to the modifications which they demand of the hypothesis that degenerating axon bundles in the goldfish brain are in some way attractive to regenerating axons.