Pretectal Nucleus Lentiformis Mesencephali Research Articles

Cerebellar Inputs in the American Alligator (Alligator mississippiensis)

Crocodilians (alligators, crocodiles, and gharials) are the closet living relatives to birds and, as such, represent a key clade to understand the evolution of the avian brain. However, many aspects of crocodilian neurobiology remain unknown. In this paper, we address an important knowledge gap as there are no published studies of cerebellar connections in any crocodilian species. We used injections of retrograde tracers into the cerebellum of the American alligator (Alligator mississippiensis) to describe for the first time the origin of climbing and mossy fiber inputs. We found that inputs to the cerebellum in the American alligator are similar to those of other nonavian reptiles and birds. Retrograde labeled cells were found in the spinal cord, inferior olive, reticular formation, vestibular and cerebellar nuclei, as well as in nucleus ruber and surrounding tegmentum. Additionally, we found no retrogradely labeled cells in the anterior rhombencephalon which suggest that, like other nonavian reptiles, crocodilians may lack pontine nuclei. Similar to birds and other nonavian reptiles, we found inputs to the cerebellum from the pretectal nucleus lentiformis mesencephali. Additionally, we found retrogradely labeled neurons in two nuclei in the pretectum: the nucleus circularis and the interstitial nucleus of the posterior commissure. These pretectal projections have not been described in any other nonavian reptile to date, but they do resemble projections from the nucleus spiriformis medialis of birds. Our results show that many inputs to the cerebellum are highly conserved among sauropsids and that extensive pretectal inputs to the cerebellum are not exclusive to the avian brain. Finally, we suggest that the pontine nuclei of birds are an evolutionary novelty that may have evolved after the last common ancestor between birds and crocodilians, and may represent an intriguing case of convergent evolution with mammals.

Pretecto‐ and ponto‐cerebellar pathways to the pigeon oculomotor cerebellum follow a zonal organization

Both birds and mammals have relatively large forebrains and cerebella. In mammals, there are extensive sensory-motor projections to the cerebellum through the pontine nuclei originating from several parts of the cerebral cortex. Similar forebrain-to-cerebellum pathways exist in birds, but the organization of this circuitry has not been studied extensively. Birds have two nuclei at the base of the brainstem that are thought to be homologous to the pontine nuclei of mammals, the medial and lateral pontine nuclei (PM, PL). Additionally, birds are unique in that they have a pretectal nucleus called the medial spiriform nucleus (SpM) that, like the pontine nuclei, also receives projections from the forebrain and projects to the oculomotor cerebellum (OCb; folia VI to VIII). The OCb also receives input from the pretectal nucleus lentiformis mesencephali (LM), which analyzes visual optic flow information resulting from self-movement. In this study, we used single or double injections of fluorescent tracers to study the organization of these inputs from PM, PL, SpM and LM to the OCb in pigeons. We found that these inputs follow a zonal organization. The most medial zone in the OCb, zone A1, receives bilateral inputs from the lateral SpM, PL and LM. Zones A2 and C receive a bilateral projection from the medial SpM, and a mostly contralateral projection from PM and LM. We discuss how the pathway to zone A1 processes mainly visuo-motor information to spinal premotor areas, whereas the pathways to zone A2/C processes somato-motor and visuo-motor information and may have a feedback/modulatory role.

Visual-Cerebellar Pathways and Their Roles in the Control of Avian Flight.

In this paper, we review the connections and physiology of visual pathways to the cerebellum in birds and consider their role in flight. We emphasize that there are two visual pathways to the cerebellum. One is to the vestibulocerebellum (folia IXcd and X) that originates from two retinal-recipient nuclei that process optic flow: the nucleus of the basal optic root (nBOR) and the pretectal nucleus lentiformis mesencephali (LM). The second is to the oculomotor cerebellum (folia VI-VIII), which receives optic flow information, mainly from LM, but also local visual motion information from the optic tectum, and other visual information from the ventral lateral geniculate nucleus (Glv). The tectum, LM and Glv are all intimately connected with the pontine nuclei, which also project to the oculomotor cerebellum. We believe this rich integration of visual information in the cerebellum is important for analyzing motion parallax that occurs during flight. Finally, we extend upon a suggestion by Ibbotson (2017) that the hypertrophy that is observed in LM in hummingbirds might be due to an increase in the processing demands associated with the pathway to the oculomotor cerebellum as they fly through a cluttered environment while feeding.

Motion parallax processing in pigeon (Columba livia) pretectal neurons

In the visual system of invertebrates and vertebrates there are specialised groups of motion-sensitive neurons, with large receptive fields, which are optimally tuned to respond to optic flow produced by the animals' movement through the 3-D world. From their response characteristics, shared frame of reference with the vestibular or inertial system, and anatomical connections, these neurons have been implicated in the stabilisation of retinal images, the control of posture and balance, and the animal's motion trajectories through the world. Using standard electrophysiological techniques and computer-generated stimuli, we show that some of these flow-field neurons in the pretectal nucleus lentiformis mesencephali in pigeons appear to be processing motion parallax. Two large overlapping planes of random dots moving independently were used to simulate motion parallax, in which one with larger dots was moved fast and the other with smaller dots was moved slowly in the opposite direction. Their neural responses to these two superimposed planes were facilitated above those produced by a single plane of moving dots and those produced by two layers moving in the same direction. Furthermore, some of these neurons preferred backward motion in the visual field and others preferred forward motion, suggesting that they may separately code visual objects 'nearer' and 'farther' than the stabilised ('on') plane during forward translational motion. A simple system is proposed whereby the relative activity in 'near', 'far' and 'on' populations could code depth through motion parallax in a metameric manner similar to that employed to code color vision and stereopsis.

Looming responses of telencephalic neurons in the pigeon are modulated by optic flow

The movement of animals through space filled with various objects requires the interaction between neuronal mechanisms specialized for processing local object motion and those specialized for processing optic flow generated by self-motion of the animal. In the avian brain, visual nuclei in the tectofugal pathway are primarily involved in the detection of object motion. By contrast, the nucleus of the basal optic root (nBOR) and the pretectal nucleus lentiformis mesencephali (nLM) are dedicated to the analysis of optic flow. But little is known about how these two systems interact. Using single-unit recording in the entopallium of the tectofugal pathway, we show that some neurons appeared to be integrating visual information of looming objects and whole-field motion simulating optic flow. They specifically responded to looming objects, but their looming responses were modulated by optic flow. Optic flow in the nasotemporal direction, typically produced by the forward movement of the bird, only mildly inhibited the looming responses. Furthermore, these neurons started firing later than when the looming object was presented against a stationary background. However, optic flow in other directions, especially the temporonasal direction, strongly inhibited their looming responses. Previous studies have implicated looming-sensitive neurons in predator avoidance behavior and these results suggest that a bird in motion may need less time to initiate an avoidance response to an approaching object.

Expression of calcium-binding proteins in cerebellar- and inferior olivary-projecting neurons in the nucleus lentiformis mesencephali of pigeons

In the avian brain, the optokinetic response is controlled by two retinal-recipient nuclei: the nucleus of the basal optic root (nBOR) of the accessory optic system and the pretectal nucleus lentiformis mesencephali (LM). Although considered sister nuclei because of their similar response properties and function, there are both similarities and differences with respect to efferent projections and neurochemistry. Both nBOR and LM project to the cerebellum (Cb) directly as mossy fibers but also indirectly via the inferior olive (IO). In a previous report, we showed that the cerebellar- and inferior olivary-projecting neurons in nBOR of pigeons differentially express the calcium-binding proteins calretinin (CR) and parvalbumin (PV). Both CR and PV are expressed in the somata of LM neurons, although the latter is not as prevalent, and whether expression of CR and PV reflects cerebellar and IO projections is not known. In this report, by combining retrograde neuronal tracing from the Cb and IO with fluorescent immunohistochemistry, we examined the expression of these calcium-binding proteins in the pigeon LM. Half (52%) of the cerebellar-projecting neurons were CR+ve, but only 15% were PV+ve. Almost all (>95%) these PV+ve cells also expressed CR. In contrast, few of the IO-projecting neurons expressed CR or PV (<or=5%). This is strikingly similar to what we observed in nBOR and reveals that calcium-binding protein expression is concordant with projection patterns in two nuclei that share similar functions.

Novel, continuous visual motion induces c-fos expression in the avian optokinetic nuclei and optic tectum

We studied the stimulus characteristics necessary for the expression of c- fos protein in optokinetic system neurons using immunocytochemistry. Using whole-field visual motion as a stimulus, we found substantial c- fos expression in the optic tectum (TeO), the nucleus of the basal optic root (nBOR) and the pretectal nucleus lentiformis mesencephali (LM); in all cases immunostaining was seen only on the side contralateral to the eye viewing whole-field unidirectional motion; the side of the brain contralateral to the eye wearing a diffuser showed no staining. In the nBOR and the LM, different regions showed a remarkable specificity of c- fos expression depending on the direction of visual motion stimulation. Neurons were stained primarily in regions known from previous electrophysiological recordings to be maximally responsive to that direction of motion; little staining was seen after motion orthogonal to the preferred motion direction. Novel, continuous visual motion stimuli, lasting more than 30 min, was required for maximal c- fos expression, suggesting that brief periods of unidirectional optic flow, as would be experienced during normal life, do not stimulate the expression of c- fos. The largest number of neurons was labeled when birds raised from hatching with one eye covered by a diffuser were exposed to full-field visual motion immediately after the diffuser was switched from one eye to the other, so that only the previously naive eye was visually stimulated. We conclude that the expression of c- fos in the optokinetic nuclei is linked to near peak firing rates on the one hand, and the novelty and duration of the visual signals, on the other, supporting the assumption that this expression is mainly related to stimulus contexts leading to neuronal plastic changes.

Corollary discharge circuits for saccadic modulation of the pigeon visual system

A saccadic eye movement causes a variety of transient perceptual sequelae that might be the results of corollary discharge. Here we describe the neural circuits for saccadic corollary discharge that modulates activity throughout the pigeon visual system. Saccades in pigeons caused inhibition that was mediated by corollary discharge followed by enhancement of firing activity in the telencephalic hyperpallium, visual thalamus and pretectal nucleus lentiformis mesencephali (nLM) with opposite responses in the accessory optic nucleus (nBOR). Inactivation of thalamic neurons eliminated saccadic responses in telencephalic neurons, and inactivation of both the nLM and the nBOR abolished saccadic responses in thalamic neurons. Saccade-related omnipause neurons in the brainstem raphe complex inhibited the nBOR and excited the nLM, whereas inactivation of raphe neurons eliminated saccadic responses in both optokinetic and thalamic neurons. It seems that saccadic responses in telencephalic neurons are generated by corollary discharge signals from brainstem neurons that are transmitted through optokinetic and thalamic neurons. These signals might have important roles in visual perception.

Projections of the nucleus of the basal optic root in pigeons (Columba livia): A comparison of the morphology and distribution of neurons with different efferent projections

The avian nucleus of the basal optic root (nBOR) is a visual structure involved in the optokinetic response. nBOR consists of several morphologically distinct cell types, and in the present study, we sought to determine if these different cell types had differential projections. Using retrograde tracers, we examined the morphology and distribution of nBOR neurons projecting to the vestibulocerebellum (VbC), inferior olive (IO), dorsal thalamus, the pretectal nucleus lentiformis mesencephali (LM), the contralateral nBOR, the oculomotor complex (OMC) and a group of structures along the midline of the mesencephalon. The retrogradely labeled neurons fell into two broad categories: large neurons, most of which were multipolar rather than fusiform and small neurons, which were either fusiform or multipolar. From injections into the IO, LM, contralateral nBOR, and structures along the midline-mesencephalon small nBOR neurons were labeled. Although there were no differences with respect to the size of the labeled neurons from these injections, there were some differences with the respect to the distribution of labeled neurons and the proportion of multipolar vs. fusiform neurons. From injections into the VbC, the large multipolar cells were labeled throughout nBOR. The only other cases in which these large neurons were labeled were contralateral OMC injections. To investigate if single neurons project to multiple targets we used paired injections of red and green fluorescent retrograde tracers into different targets. Double-labeled neurons were never observed indicating that nBOR neurons do not project to multiple targets. We conclude that individual nBOR neurons have unique projections, which may have differential roles in processing optic flow and controlling the optokinetic response.

Neural specialization for hovering in hummingbirds: Hypertrophy of the pretectal nucleus lentiformis mesencephali

Hummingbirds possess an array of morphological and physiological specializations that allow them hover such that they maintain a stable position in space for extended periods. Among birds, this sustained hovering is unique to hummingbirds, but possible neural specializations underlying this behavior have not been investigated. The optokinetic response (OKR) is one of several behaviors that facilitates stabilization. In birds, the OKR is generated by the nucleus of the basal optic root (nBOR) and pretectal nucleus lentiformis mesencephali (LM). Because stabilization during hovering is dependent on the OKR, we predicted that nBOR and LM would be significantly enlarged in hummingbirds. We examined the relative size of nBOR, LM, and other visual nuclei of 37 species of birds from 13 orders, including nine hummingbird species. Also included were three species that hover for short periods of time (transient hoverers; a kingfisher, a kestrel, and a nectarivorous songbird). Our results demonstrate that, relative to brain volume, LM is significantly hypertrophied in hummingbirds compared with other birds. In the transient hoverers, there is a moderate enlargement of the LM, but not to the extent found in the hummingbirds. The same degree of hypertrophy is not, however, present in nBOR or the other visual nuclei measured: nucleus geniculatus lateralis, pars ventralis, and optic tectum. This selective hypertrophy of LM and not other visual nuclei suggests that the direction-selective optokinetic neurons in LM are critical for sustained hovering flight because of their prominent role in the OKR and gaze stabilization.

Two optic flow pathways from the pretectal nucleus lentiformis mesencephali to the cerebellum in pigeons (Columba livia)

Neurons in the pretectal nucleus lentiformis mesencephali (LM) are involved in the analysis of optic flow. LM provides mossy fiber inputs to folia VI-VIII of the posterior cerebellum and IXcd of the vestibulocerebellum. Previous research has shown that the vestibulocerebellum is involved in visual-vestibular integration supporting gaze stabilization. The function of folia VI-VIII in pigeons is not well understood; however, these folia receive input from a tectopontine system, which is likely involved with analyzing local motion as opposed to optic flow. We sought to determine whether the mossy fiber input from LM to IXcd differs from that to VI-VIII. Fluorescent retrograde tracers were injected into these folia, and the pattern of labeling in LM was observed. Large multipolar neurons were labeled throughout the rostrocaudal extent of LM. There was a clear mediolateral difference: 74.3% of LM neurons projecting to IXcd were located in the lateral subnucleus of LM (LMl), whereas 73.8% of LM neurons projecting to VI-VIII were found in medial LM (LMm). This suggests that the subnuclei of LM have differing roles. In particular, the LMl-IXcd pathway is involved in generating the optokinetic response. We suggest that the pathway from LMm to VI-VIII is integrating optic flow and local motion to support various oculomotor and visuomotor behaviors, including obstacle avoidance during locomotion.

Differential modulation of thalamic neurons by optokinetic nuclei in the pigeon

The visual system in the pigeon is composed of the tectofugal, thalamofugal and accessory optic pathways. Though their anatomy and physiology have been extensively studied, the functional interactions between these pathways are largely unknown. The present study shows by using multiple electrophysiological techniques that firing activity in the nucleus opticus principalis thalami (OPT) of the thalamofugal pathway is differentially modulated by the pretectal nucleus lentiformis mesencephali (nLM) and the nucleus of the basal optic root (nBOR) of the accessory optic system, two optokinetic nuclei responsible for generating eye movements to stabilize the image on the retina. Reversible inactivation, electrical stimulation, microiontophoresis and receptive field mapping experiments all consistently indicate that the nBOR-OPT pathway is inhibitory and mediated by GABA as a transmitter and its GABA A receptors, whereas the nLM-OPT pathway is excitatory and mediated by glutamate as a transmitter and its NMDA receptors. They also differentially modulate the size and/or responsiveness of receptive fields in OPT cells as well. Numerous electrode tip sites were histologically confirmed in the neural structures under study. The results suggest that these optokinetic nuclei may dually modulate the transfer of visual information from the retina to the telencephalon at the thalamic level during eye movements.

Telencephalic projections to the nucleus of the basal optic root and pretectal nucleus lentiformis mesencephali in pigeons

In birds, the nucleus of the basal optic root (nBOR) of the accessory optic system (AOS) and the pretectal nucleus lentiformis mesencephali (LM) are involved in the analysis of optic flow and the generation of the optokinetic response. In several species, it has been shown that the AOS and pretectum receive input from visual areas of the telencephalon. Previous studies in pigeons using anterograde tracers have shown that both nBOR and LM receive input from the visual Wulst, the putative homolog of mammalian primary visual cortex. In the present study, we used retrograde and anterograde tracing techniques to further characterize these projections in pigeons. After injections of the retrograde tracer cholera toxin subunit B (CTB) into either LM or nBOR, retrograde labeling in the telencephalon was restricted to the hyperpallium apicale (HA) of the Wulst. From the LM injections, retrograde labeling appeared as a discrete band of cells restricted to the lateral edge of HA. From the nBOR injections, the retrograde labeling was more distributed in HA, generally dorsal and dorso-medial to the LM-projecting neurons. In the anterograde experiments, biotinylated dextran amine (BDA) was injected into HA and individual axons were reconstructed to terminal fields in the LM and nBOR. Those fibers projecting to the nBOR also innervated the adjacent ventral tegmental area. However, tracing of BDA-labeled axons revealed no evidence that individual neurons project to both LM and nBOR. In summary, our results suggest that the nBOR and LM receive input from different areas of the Wulst. We discuss how these projections may transmit visual and/or somatosensory information to the nBOR and LM.

The Accessory Optic System Contributes to the Spatio-Temporal Tuning of Motion-Sensitive Pretectal Neurons

The nucleus of the basal optic root (nBOR) of the accessory optic system (AOS) and the pretectal nucleus lentiformis mesencephali (LM) are involved in the analysis of optic flow that results from self-motion and are important for oculomotor control. These neurons have large receptive fields and exhibit direction selectivity to large moving stimuli. In response to drifting sine wave gratings, LM and nBOR neurons are tuned to either low spatial/high temporal frequencies (SF, TF) or high SF/low TF stimuli. Given that velocity = TF/SF, these are referred to as "fast" and "slow" neurons, respectively. There is a heavy projection from the AOS to the pretectum, although its function is unknown. We recorded the directional and spatio-temporal tuning of LM units in pigeons before and after nBOR was inactivated by tetrodotoxin injection. After nBOR inactivation, changes in direction preference were observed for only one of 18 LM units. In contrast, the spatio-temporal tuning of LM units was dramatically altered by nBOR inactivation. Two major effects were observed. First, in response to motion in the preferred direction, most (82%) neurons showed a substantially reduced (mu = -67%) excitation to low SF/high TF gratings. Second, in response to motion in the anti-preferred direction, most (63%) neurons showed a dramatically reduced (mu = -78%) inhibition to high SF/low TF gratings. Thus the projection from the nBOR contributes to the spatio-temporal tuning rather than the directional tuning of LM neurons. We propose a descriptive model whereby LM receives inhibitory and excitatory input from "slow" and "fast" nBOR neurons, respectively.

Fast and slow neurons in the nucleus of the basal optic root in pigeons

The nucleus of the basal optic root (nBOR) is involved in the generation of the optokinetic response. Previous studies showed that most nBOR neurons exhibit direction-selectivity in response to largefield motion. We investigated the responses of pigeon nBOR neurons to drifting sine wave gratings of varying spatial and temporal frequency (SF,TF). Two groups of neurons were revealed. The first group preferred gratings of low SF (mean, 0.07 cycles per degree (cpd)) and high TF (mean, 0.76 Hz) (‘fast’ stimuli). The second group preferred gratings of high SF (mean, 0.56 cpd) and lower TF (mean, 0.33 Hz) (‘slow’ stimuli). Previous studies have demonstrated fast and slow neurons in pretectal nucleus lentiformis mesencephali, which is also involved in the generation of the optokinetic response.

Spatiotemporal properties of fast and slow neurons in the pretectal nucleus lentiformis mesencephali in pigeons.

Neurons in the pretectal nucleus lentiformis mesencephali (LM) are involved in the analysis of optic flow that results from self-motion. Previous studies have shown that LM neurons have large receptive fields in the contralateral eye, are excited in response to largefield stimuli moving in a particular (preferred) direction, and are inhibited in response to motion in the opposite (anti-preferred) direction. We investigated the responses of LM neurons to sine wave gratings of varying spatial and temporal frequency drifting in the preferred and anti-preferred directions. The LM neurons fell into two categories. "Fast" neurons were maximally excited by gratings of low spatial [0.03-0.25 cycles/ degrees (cpd)] and mid-high temporal frequencies (0.5-16 Hz). "Slow" neurons were maximally excited by gratings of high spatial (0.35-2 cpd) and low-mid temporal frequencies (0.125-2 Hz). Of the slow neurons, all but one preferred forward (temporal to nasal) motion. The fast group included neurons that preferred forward, backward, upward, and downward motion. For most cells (81%), the spatial and temporal frequency that elicited maximal excitation to motion in the preferred direction did not coincide with the spatial and temporal frequency that elicited maximal inhibition to gratings moving in the anti-preferred direction. With respect to motion in the anti-preferred direction, a substantial proportion of the LM neurons (32%) showed bi-directional responses. That is, the spatiotemporal plots contained domains of excitation in addition to the region of inhibition. Neurons tuned to stimulus velocity across different spatial frequency were rare (5%), but some neurons (39%) were tuned to temporal frequency. These results are discussed in relation to previous studies of the responses of neurons in the accessory optic system and pretectum to drifting gratings and other largefield stimuli.

Binocular neurons in the nucleus lentiformis mesencephali in pigeons: responses to translational and rotational optic flowfields

The pretectal nucleus lentiformis mesencephali (LM) receives direct input from the contralateral retina and is dedicated to the analysis of optic flowfields resulting from self-motion. The activity of 126 LM neurons in response to optic flow stimuli was recorded. As with previous studies, it was found that most neurons (∼90%) exhibited direction-selectivity to large-field stimuli moving in the contralateral hemifield. However, some neurons (∼10%) responded to stimulation of both eyes and had receptive field structures conducive for detection of particular patterns of optic flow resulting from either self-translation or self-rotation. These binocular neurons were maximally responsive to panoramic optic flowfields simulating either translational or rotational optic flow.

Projections from the nucleus of the basal optic root and nucleus lentiformis mesencephali to the inferior olive in pigeons (Columba livia)

The nucleus of the basal optic root (nBOR) of the accessory optic system (AOS) and the pretectal nucleus lentiformis mesencephali (LM) are involved in the analysis of optic flow and the generation of the optokinetic response. Previous studies have shown that the nBOR projects bilaterally to the medial column (mc) of the inferior olive (IO) and the LM projects to the ipsilateral mc. In the present study the retrograde tracer cholera toxin subunit B was injected into either the caudal or rostral mc. From all injections, retrogradely labeled cells were seen in the ipsilateral pretectum along the border of the medial and lateral subnuclei of the LM. Cells were also seen in bilaterally in the nBOR. On the contralateral side, a discrete group of cells was labeled in the rostral margin of the nBOR. These cells were localized in the dorsal portion of the nBOR proper and some were found in the adjacent nBOR dorsalis. On the ipsilateral side, a diffuse group of cells was seen in the caudal nBOR. Most of these cells were in the nBOR dorsalis and outside the nBOR complex in the area ventralis of Tsai and the reticular formation. From the injections into the caudal mc, a greater proportion of labeled cells was found in the LM, whereas a greater proportion of cells was found in the nBOR from the injections into the rostral mc. This differential projection from LM and nBOR to the caudal and rostral mc is consistent with the optic flow preferences of neurons in the mc, and a similar pattern of connectivity has been found in mammalian species.

The pigeon optokinetic system: visual input in extraocular muscle coordinates.

The generation of compensatory eye movements in response to rotational head movements involves the transformation of visual-optokinetic and vestibular signals into commands controlling the appropriate eye muscles. Previously, it has been shown that the three systems (optokinetic, vestibular, and eye muscle) share a similar three-dimensional reference frame. In this report, we suggest that a peculiarity in the structure of the horizontal recti in pigeons demonstrates that the optokinetic system is organized with respect to the eye muscles rather than the vestibular canals. Measurements of the orientation of the plane for each of the lateral and medial recti were obtained. These were compared with the direction preferences of optokinetic neurons responsive to horizontal motion, namely "back" units in the nucleus of the basal optic root (nBOR), "forward" units in the pretectal nucleus lentiformis mesencephali (LM), and "vertical axis" (VA) Purkinje cells in the flocculus. The average direction preference of LM neurons excited in response to forward (temporal to nasal) visual motion, and VA Purkinje cells in response to optokinetic motion in the ipsilateral visual field was approximately parallel to the visual horizontal. This corresponded to the orientation of the medial rectus, which was also approximately parallel to the visual horizontal. The average direction preference of nBOR neurons excited in response to backward (nasal to temporal) visual motion, and VA Purkinje cells in response to optokinetic motion in the contralateral visual field was approximately 20-30 deg down from the visual horizontal. The orientation of the lateral rectus was also approximately 20-30 deg down from the visual horizontal. These data suggest that the incoming optokinetic signals are organized with respect to the outgoing extraocular muscle commands.

Distribution of GAD-like immunoreactivity in the retina and central visual system of Rana pipiens.

Immunohistochemistry was used to survey the distribution of glutamic acid decarboxylase (GAD), the synthesizing enzyme for gamma-aminobutyric acid (GABA), throughout the visual system of the frog Rana pipiens. GAD-like immunoreactivity (GAD-LI) was assessed in the retina, in primary retinorecipient targets, and in thalamic nuclear groups postsynaptic to primary retinal terminal zones. Within the retina, the inner plexiform layer displayed intense GAD-LI, but immunoreactivity was absent in the ganglion cell layer (GCL). Putative amacrine, bipolar, and horizontal cell somata were also labeled. Centrally, GAD-LI was observed in all primary visual nuclei. Dense, GAD-like immunoreactive punctate structures (puncta), presumably terminals, were observed in the pretectal nucleus lentiformis mesencephali, posterior thalamic neuropil, and uncinate neuropil. GAD-like immunoreactive puncta were noted in several laminae of the optic tectum, with the highest concentrations located within the 9th and 8th laminae. Moderate numbers of GAD-like immunoreactive puncta were found in the mesencephalic nucleus of the basal optic root, and two thalamic neuropils--corpus geniculatum and neuropil of Bellonci. The ventrolateral area, posterocentral nucleus, and posterolateral nucleus all contained sparse amounts of GAD-LI. These observations suggest that GABA plays an important physiological role in all central visual areas of Rana pipiens.