The centrifugal visual system of a palaeognathous bird,the Chilean Tinamou (Nothoprocta perdicaria)

The avian centrifugal visual system, which projects from the brain to the retina, has been intensively studied in several Neognathous birds that have a distinct isthmo‐optic nucleus (ION). However, birds of the order Palaeognathae seem to lack a proper ION in histologically stained brain sections. We had previously reported in the palaeognathous Chilean Tinamou (Nothoprocta perdicaria) that intraocular injections of Cholera Toxin B subunit retrogradely label a considerable number of neurons, which form a diffuse isthmo‐optic complex (IOC). In order to better understand how this IOC‐based centrifugal visual system is organized, we have studied its major components by means of in vivo and in vitro tracing experiments. Our results show that the IOC, though structurally less organized than an ION, possesses a dense core region consisting of multipolar neurons. It receives afferents from neurons in L10a of the optic tectum, which are distributed with a wider interneuronal spacing than in Neognathae. The tecto‐IOC terminals are delicate and divergent, unlike the prominent convergent tecto‐ION terminals in Neognathae. The centrifugal IOC terminals in the retina are exclusively divergent, resembling the terminals from “ectopic” centrifugal neurons in Neognathae. We conclude that the Tinamou's IOC participates in a comparable general IOC‐retina‐TeO‐IOC circuitry as the neognathous ION. However, the connections between the components are structurally different and their divergent character suggests a lower spatial resolution. Our findings call for further comparative studies in a broad range of species for advancing our understanding of the evolution, plasticity and functional roles of the avian centrifugal visual system.     

Functional anatomy of the avian centrifugal visual system

Although first described over a century ago, the centrifugal visual system (CVS) projecting to the retina still remains somewhat of an enigma with regard to its functional role in visually-guided behavior. The highly developed avian CVS has been the most extensively investigated and the anatomical organization of its two component centrifugal structures, the n. isthmo-opticus (NIO) and ectopic neurons (EN), including its afferent brainstem projections is reviewed. The results of double-labeling studies combining axonal tracing techniques and immunohistofluorescence have demonstrated GABA immunoreactivity (-ir) of interneurons within the neuropilar zone of the NIO, choline acetyltransferase (ChAT)-ir and nitric oxide synthase (NOS)-ir in the centrifugal cells of the NIO and EN as well as in the afferent projection neurons of layers 9/10 of the optic tectum. The data are discussed in terms of neurochemical and excitatory/inhibitory mechanisms within the different components of the avian CVS in relation to hypotheses which have implicated this system in visual attention and ground-feeding behavior.     

Visual afferents from an eye in the terrestrial slug Limax valentianus

Terrestrial gastropods have a lens-bearing eye on the tip of their tentacles. There are two morphologically distinct photoreceptors, called Type-I and Type-II photoreceptors, in the retina. Type-I photoreceptors are equipped with highly developed photoreceptive microvilli in their outer rhabdomeric segment, whereas Type-II photoreceptors have short and fewer microvilli. Although both types of photoreceptors send afferent projections directly to the brain, their destinations in the brain, called optic neuropiles, have not been sufficiently investigated. Our recent studies revealed that there are commissural fibers in the cerebral ganglia that transmit photic information acquired by bilateral eyes. Moreover, some of the retinal photoreceptors are connected by gap junctions to the photosensitive brain neurons, suggesting the functional interaction of the photic information between the eye and brain photoreceptors, as well as between bilateral eyes. However, it has not been clarified which type of retinal photoreceptors send commissural projections to the contralateral hemiganglion nor interact with the brain photoreceptors. In the present study, we demonstrated by molecular histological analyses and tracer injections that (1) Type-I and Type-II photoreceptors send glutamatergic afferent projections to the medial and lateral lobes of the ipsilateral optic neuropile, respectively, (2) direct synaptic interaction between bilateral optic nerves occurs in the medial lobe of the optic neuropile, and (3) brain photosensory neurons form gap junctions with the medial lobe of the contralateral optic neuropile. These results reveal an ordered pattern of afferent projections from the retina and provide insight into the different functional roles of retinal photoreceptors.     

Reorganization of visual pathways following posthatching removal of one retina in pigeons

Organization of visual pathways was studied in 2-month-old pigeons that underwent unilateral retinal removal on either the day of hatching (ERA, i.e., early retinal ablated) or the 9th day after hatching (LRA, i.e., late retinal ablated). A general size reduction of visual areas contralateral to the removed retina was found in ERA pigeons, which additionally showed an altered differentiation of thalamic visual targets as well as a different cytoarchitectonic arrangement of the superficial layers of the optic tectum. No comparable modifications were found in LRA pigeons. The retinal projections of the remaining eye were studied following intraocular injections of 3H-proline. Both in ERA and LRA pigeons, the distribution of retinofugal afferents to primary visual regions contralateral to the injected eye was similar to that of control pigeons. Anomalous ipsilateral projections from the remaining retina to primary retinorecipient regions were found in ERA pigeons only. Effects of early ablation of one retina on second-order visual connections were also studied. Following injections of wheat germ agglutinin-conjugated horseradish peroxidase (WGA-HRP) into the visual Wulst contralateral to the operated eye, a smaller number of ipsilateral projecting thalamo-Wulst neurons was found as compared with control pigeons. In contrast, the contralateral thalamo-Wulst projections were increased. No changes in thalamo-Wulst projections were found following tracer injections into the opposite Wulst, i.e., ipsilateral to the operated eye. The present study demonstrates a substantial anatomical reorganization of both primary and secondary visual pathways following unilateral retinal removal immediately after hatching, when maturation of the visual system is not yet completed.     

Visual system of a naturally microphthalmic mammal: The blind mole rat,Spalax ehrenbergi

Retinal projections and visual thalamo-cortical connections were studied in the subterranean mole rat, belonging to the superspecies Spalax ehrenbergi, by anterograde and retrograde tracing techniques. Quantitative image analysis was used to estimate the relative density and distribution of retinal input to different primary visual nuclei. The visual system of Spalax presents a mosaic of both regressive and progressive morphological features. Following intraocular injections of horseradish peroxidase conjugates, the retina was found to project bilaterally to all visual structures described as receiving retinal afferents in non-fossorial rodents. Structures involved in form analysis and visually guided behaviors are reduced in size by more than 90%, receive a sparse retinal innervation, and are cytoarchitecturally poorly differentiated. The dorsal lateral geniculate nucleus, as defined by cyto- and myelo-architecture, cytochrome oxidase, and acetylcholinesterase distribution as well as by afferent and efferent connections, consists of a narrow sheet 3–5 neurons thick, in the dorsal thalamus. Connections with visual cortex are topographically organized but multiple cortical injections result in widespread and overlapping distributions of geniculate neurons, thus indicating that the cortical map of visual space is imprecise. The superficial layers of the superior colliculus are collapsed to a single layer, and the diffuse ipsilateral distribution of retinal afferents also suggests a lack of precise retinotopic relations. In the pretectum, both the olivary pretectal nucleus and the nucleus of the optic tract could be identified as receiving ipsilateral and contralateral retinal projections. The ventral lateral geniculate nucleus is also bilaterally innervated, but distinct subdivisions of this nucleus or the intergeniculate leaflet could not be distinguished. The retina sends a sparse projection to the dorsal and lateral terminal nuclei of the accessory optic system. The medial terminal nucleus is not present. In contrast to the above, structures of the “non-image forming” visual pathway involved in photoperiodic perception are well developed in Spalax. The suprachiasmatic nucleus receives a bilateral projection from the retina and the absolute size, cytoarchitecture, density, and distribution of retinal afferents in Spalax are comparable with those of other rodents. A relatively hypertrophied retinal projection is observed in the bed nucleus of the stria terminalis. Other regions which receive sparse visual input include the lateral and anterior hypothalamic areas, the retrochiasmatic region, the sub-paraventricular zone, the paraventricular hypothalamic nucleus, the anteroventral and anterodorsal nuclei, the lateral habenula, the mediodorsal nucleus, and the basal telencephalon. These results indicate that the apparently global morphological regression of the visual system conceals a selective expansion of structures related to functions of photoperiodic perception and photo-neuroendocrine regulation. We suggest that the evolution of an atrophied eye and reduced visual system is an adaptively advantageous response to the unique subterranean environment. Factors favoring regression include mechanical aspects, metabolic constraints, and competition between sensory systems. The primary advantage of sensory atrophy is the metabolic economy gained by the reduction of visual structures which do not contribute significantly to the animal's fitness. © 1993 Wiley-Liss, Inc.     

Dendrite development and target innervation of displaced retinal ganglion cells of the chick (Gallus gallus)

The avian accessory optic system (AOS) processes visual signals of translational and rotational flowfields resulting from self-motion. It has been investigated extensively with physiological methods and, because of its anatomical distinction from other retinofugal projections, is well suited for the investigation of dendritic differentiation and axonal pathfinding. Displaced retinal ganglion cells (dRGC) constitute the retinal origin of the AOS. Since little is known about the time course of the development of this projection, we studied the dendritic differentiation of dRGC, their innervation of the nucleus of the basal optic root (nBOR) and the histological development of this target area. dRGC, visualized by retrograde 1,1'-dioctadecyl-3,3,3',3'-tetramethyl-indocarbocyanine perchlorate labeling, migrated into the inner nuclear layer of the retina and subsequently developed their characteristic dendritic morphology between E9 and E14. At this stage, dendrites were unistratified in the inner plexiform layer and displayed characteristic branches with 45-90 degrees angles. The frequency of dendritic branches increased from an average of 44 branches per cell at E9 to an average of 155 at E15. This phase was followed by a period of dendritic pruning, E15-E17, where a large number of small branches were eliminated. At the time of hatching, dRGC were morphologically mature with mean dendritic field sizes of 0.28 mm2 and an average of 108 dendritic branches per cell. Retinal innervation of the nBOR occurred between E8 and E11, and tracer injections at later stages revealed no further changes. In addition to the predominant contralateral projection, we have also described a connection to the ipsilateral nBOR. This ipsilateral pathway persisted at least to juvenile stages (P14). The histological development of the nBOR proceeded such that calretinin-immunoreactive neurons were observed from E10 onwards and morphologically described cell types evolved after E12.     

The retinohypothalamic tract originates from a distinct subset of retinal ganglion cells

The retinal ganglion cells giving rise to retinohypothalamic projections in the rat were identified using retrograde transport of horseradish peroxidase (HRP) or Fluoro Gold injected into the suprachiasmatic nucleus (SCN), and using transneuronal transport of the Bartha strain of the swine herpesvirus (PRV-Bartha). When PRV-Bartha is injected into one eye, it is taken up by retinal ganglion cells, replicated, transported to axon terminals in the SCN, and released. There the virus may take one, or both, of two paths to retinal ganglion cells in the contralateral eye: (1) uptake by SCN neurons, replication, and release from the neurons with uptake and retrograde transport in retinal afferents originating in the contralateral retina; (2) transneuronal passage through axo-axonic appositions between retinal afferents in the SCN with subsequent retrograde transport of virus to the contralateral retina. The ganglion cells thus labeled are a homogeneous population of small neurons (mean diameter, 12.8 ± 2.2 μm an mean area, 81.8 ± 21.8 μm²) with sparsely branching dendrites that are widely distributed over the retina. This population is best identified when virus labeling of retinal projections in areas beyond the hypothalamus is eliminated by lateral geniculate lesions that transect the optic tract at its entry into the geniculate complex. The same population is labeled with retrograde tracers but, with both HRP and Fluoro Gold, other ganglion cells are labeled, presumably from uptake by fibers of passage, indicating that the virus is a more reliable marker for ganglion cells giving rise to retinohypothalamic projections. The ganglion cells identified correspond to a subset of type III, or W, cells. © 1995 Wiley-Liss, Inc.     

The organization of the fibers in the optic nerve of normal and tectum-less Rana pipiens

We have examined the detailed order of retinal ganglion cell (RGC) axons in the optic nerve and tract of the frog, Ranapipiens. By using horseradish peroxidase (HRP) injections into small regions of theretina, the tectum, and at various points along the visual pathway, it hasbeen possible to follow labelled fibers throughout their course in the nerve and tract. Several surprising features in the order of fibers in the visual pathway were discovered in our investigation. The fascicular pattern of RGC axons in che retina is similar to that described in other vertebrates; however, immediately central to their entry into the optic nerve head, approximately half of the fibers from the nasal or temporal retina cross over to the opposite side of the nerve. Although the axons from the dorsal and ventral regions of the retina generally remain in the dorsal and ventral regions of the nerve, some fiber crossing occurs in those axons as well. The result of this seemingly complex rearrangement is that the optic nerve of Rana pipiens contains mirror symmetric representations of the retinal surface on either side of the dorsal ventral midline of the nerve. The fibers in each of these representation are arranged as semicircles representing the full circumference of the retina. This precise fiber order is preserved in the nerve until immediately periphearal to the optic chiasm, at which point age-related axon from both side of the nerve bundle together. Consequently, when a small pellet of HRP is placed in the chiasmic region of the nerve, an annualus of retinal ganglion cells and a corresponding annulus of RGC terminals in the tectum are la belled. As the age-related bundles of fibers emerge from the chiasm they split to form a medial bundle and a lateral bundle, which grow in the medial and lateral branches of the optic tract, respectively. Although the course followed by RGC axons in the visual pathv/ay is complex, we propose a model in which the organization of fibers in the nerve and tract can arise from a few rules of axon guidance. To determine whether the optic tecta, the primary retinal targets, play a role in the development and organization of the optic nerve and tract, we removed the tectal primordia in Rana embryos and examined the order in the nerve when the animals had reached larval stages. We found that the order in the nerve and tract was well preserved in tectumless frogs. Therefore, we propose that guidance factors independent of the target direct axon growth in the frog visual system.     

Synaptic transmission of excitation from the retina to cells in the pigeon's optic tectum

Intracellular recordings were used to study the synaptic excitation of optic tectum neurons in the pigeon. Electrical stimulation of both contralateral optic nerve and ipsilateral optic tract evoked in the tectal neurons EPSPs which in most cases were followed by an IPSP. An extrapolation procedure based on response latency was used to reveal that the EPSPs were mediated by way of mono-, di- and polysynaptic connections with the retinal endings. The laminar location of the recorded cells was estimated according to the field potential and the recording depth with the exception of the cell which was intracellulary stained with HRP. Monosynaptic EPSPs were recorded from cells in the retinorecipient region (sublayers IIa-f) as well as in the non-retinorecipient region (sublayers IIg-j and layer III) of the tectum, while di- and polysynaptic EPSPs were never recorded from the input layers. Tectofugal projections arise largely from layer III neurons. Thus, these results indicate that retinal excitation is transmitted to the output tectal cells by way of mono-, di- or polysynaptic pathways. The conduction velocities of most retinal fibers mediating the EPSP ranged from 4 to 22 m/s (average 12 m/s). However, in a number of retinal fibers the conduction velocities were in a faster range, up to 36 m/s.     

Retinal projections and retinal ganglion cell distribution patterns in a sturgeon (Acipenser transmontanus), a non-teleost actinopterygian fish

Retinal projections in a sturgeon were studied by injecting biocytin or HRP into the optic nerve. The target areas are the preoptic area, thalamus, area pretectalis, nucleus of posterior commissure, optic tectum, and nuclei of the accessory optic tract. Furthermore, a few labeled fibers and terminals were found in a ventrolateral area of the caudal telencephalon. All retinal projections are bilateral, although contralateral projections were more heavily labeled. Retrogradely labeled neurons were found in the ventral thalamus bilaterally. Retinal projections in sturgeons are similar to those of other non-teleost actinopterygians and chondrichthyans (sharks), in terms of the targets and extent of bilateral projections. Distribution patterns of ganglion cells in the retina were examined in Nissl-stained retinal whole mount preparations. The highest density areas were found in the temporal and nasal retinas, and a dense band of ganglion cells was observed along the horizontal axis between the nasal and temporal areas of highest density. The density of ganglion cells in the dorsal retina is the lowest. The total number of ganglion cells was estimated to be about 5 x 10(4) in a retina. The existence of a low density area in the dorsal retina suggests reduced visual acuity in the ventral visual field.     

Fate of uncrossed retinal projections following early or late prenatal monocular enucleation in the mouse

In mammals binocular vision is made possible by the existence in the temporal retina of ipsilaterally projecting ganglion cells (IGCs) (with axons that do not cross the brain midline and join optic fibers from the opposite eye). To learn whether early interactions between fibers of each eye play a role in generating a mixed ipsi + contralateral projection pattern, we studied with horseradish peroxidase the origin of uncrossed retinal projections in mice that developed after one eye was destroyed at very early embryonic ages. One eye was removed on embryonic day 16 (E16; when optic fibers have grown past the chiasm bilaterally, but very few have grown into the visual centers) or on E13 or E12 (when few or no optic fibers have passed the presumptive chiasm region). Normal adult mice have a mean of 946 IGCs (range: 784-1,073) within the temporal sector of the retina, and less than 25 in the rest of the retina. In adult mice enucleated at E16, an average of 1,354 (1,215-1,484) IGCs are present within a clearly demarcated temporal sector of the remaining retina and 265 (152-312) are present throughout the rest of the retina. In both the temporal and nasal retina the excess IGCs in these mice have, generally, very small somas. In some of these mice the most peripheral part of the temporal sector contains fewer IGCs. In E12 or E13 enucleates, IGCs are also generally located in a narrow (often narrower than normal) region along the temporo-inferior retinal border, but their number is less than in normal or E16-enucleated mice: E13 enucleates have a mean of 639 cells (range: 361-875) in the temporal sector and 109 (8-275) in the rest of the retina. Following enucleation of one mouse at E12, the respective values are 349 and 31 cells. The reduction in numbers of IGCs in these mice is especially pronounced for ganglion cells with small cell bodies. These findings suggest that the development of uncrossed projections in mice depends on selective guidance mechanisms of axons from temporal retina through the chiasm. These may consist of interactions of optic axons with guidance cues distributed in the presumptive chiasm (possibly at early stages) and also of fiber-fiber guidance mechanisms, in particular between fibers from each eye.     

Visual system degeneration in the glaucomatous albino quail

The peripheral (eye, retina, optic nerve) and central (primary optic tracti and centers, centrifugal visual tractus and nucleus) visual system of an imperfect albino quail mutant with a sex linked recessive gene was examined in 32 specimens ages 1 week - 16 months-hatch using various histological techniques. During the first weeks the visual system was normal and comparable in its overall organization to that found in the pigmented quail. However, the ipsilateral retinal projections were observed to be weaker in the young mutant, then completely disappeared two months after birth. Initial signs of the bupthalmos, a form of spontaneous glaucoma, appeared between the 3rd and 5th months. This was characterized by a distention of the eye linked to an increase in intraocular pressure. The pathological process was progressive and at 16 months the eye was very prominent, the anterior chamber deep and a large and globular cornea was noted. The glaucoma progressively induced different histopathological changes in the visual system including: cupping of the optic disc, degeneration of optic axons and their parent ganglion and centrifugal cells and cavernous degeneration. All of these phenomena were identifiable at about the 10th post-natal month and progressed in a relatively constant and orderly manner. The retinal projections to the nucleus ectomamillaris, ventral and lateral optic tectum and ventral pretectum were the first to degenerate. The degeneration of optic fibers attaining the dorsal pretectum and dorsal thalamus occurred later. Furthermore the retrograde degeneration in the centrifugal isthmo-optic nucleus progressed from the external to the internal pole. The mechanisms involved in the selective degeneration of centrifugal and centripetal optic fibers is discussed.     

Electron microscopic study of immunocytochemically labeled centrifugal fibers in the goldfish retina

The centrifugal fibers innervating the goldfish retina were studied quantitatively by light and electron microscopy. These fibers originating from cell bodies in the olfactory bulb were labeled by antiserum to the tetrapeptide Phe-Met-Arg-Phe-NH2 (FMRFamide). The number of FMRFamide-immunoreactive (ir) centrifugal fibers in each eye of the adult goldfish (body length: 12-15 cm) was 65 +/- 14 (mean +/- S.D., n = 7). All of these fibers in the optic nerve and the retina were unmyelinated. Each FMRFamide-ir centrifugal fiber runs along the optic fiber layer and gives several terminal arborizations in the outermost layer (layer 1) of the inner plexiform layer. Layer 1 is, therefore, densely covered by a plexus of terminal arborizations. Along these terminal arborizations, we found output synapses characterized by a cluster of small clear vesicles (40 nm in diameter) at the presynaptic site and a thickened membrane in the apposed retinal cell processes. In a sample area of 2,000 microns 2, such synapses occurred at a density of one per 105 microns 2, or about 13,000 per centrifugal fiber. Thus, the FMRFamide-ir centrifugal fibers are likely to modulate retinal cell activity through an estimated total of 840,000 output synapses per retina.     

Selective projection patterns from subtypes of retinal ganglion cells to tectum and pretectum: Distribution and relation to behavior

An important issue to understand is how visual information can influence the motor system and affect behavior. Using the lamprey (Petromyzon marinus) as an experimental model we examined the morphological subtypes of retinal ganglion cells and their projection pattern to the tectum, which controls eye, head, and body movements, and to the pretectum, which mediates both visual escape responses and the dorsal light response. We identified six distinct morphological types of retinal ganglion cell. Four of these distribute their dendrites in the inner plexiform layer (image forming layer) and project in a retinotopic manner to all areas of the tectum. The posterior part of the retina has the highest density of ganglion cells and projects to the rostral part of the tectum, in which the visual field in front of the lamprey will be represented. From this area both orienting and evasive behaviors can be elicited. In contrast, pretectum receives input from two ganglion cells types that send their dendrites only to the outer plexiform layer or the outer limiting membrane and therefore may directly contact photoreceptors, and transmit information without additional delay to pretectum, which may be particularly important for visual escape responses. One of these two types, the bipolar ganglion cell, is only found in a small patch of retina just ventral of the optic nerve. Due to its distribution, morphology, and projections we suggest that this cell may control the dorsal light response. J. Comp. Neurol. 517:257–275, 2009. © 2009 Wiley‐Liss, Inc.     

Early retinal development in the zebrafish,Danio rerio: Light and electron microscopic analyses

The morphological differentiation of the zebrafish retina was analyzed by using light (LM) and transmission electron (TEM) microscopy between the time of initial ganglion cell differentiation (≈32 hours postfertilization; hpf) and shortly after the point when the retina appears functional (≈74 hpf), i.e., when all major cell types and basic synaptic connections are in place. The results show that the inner retinal neurons, like the photoreceptor and ganglion cells, differentiate first within the ventronasal region, and differentiation subsequently spreads asymmetrically into the nasal and dorsal regions before reaching the ventrotemporal retina. In addition, we show that the attenuation of the optic stalk occurs in parallel with ganglion cell differentiation between 32 and 40 hpf. The first conventional synapses appear within the inner plexiform layer simultaneously with the first photoreceptor outer segment discs at 60 hpf; functional ribbon triads arise within photoreceptor synaptic terminals at 65 hpf; and synaptic ribbons occur within bipolar cell axon terminals at the time larvae exhibit their first visual responses (≈70 hpf). Although development is initially more advanced within the ventronasal region between 50 and 60 hpf, development across the retina rapidly equilibrates such that it is relatively comparable within all quadrants of the central retina by 70 hpf. An area within the temporal retina characterized by tightly packed and highly tiered cones emerges with subsequent development. Retinal differentiation in the zebrafish corresponds with that generally described in other vertebrates and can be correlated with the development of visual and electroretinographic responses in the animal. J. Comp. Neurol. 404:515–536, 1999. © 1999 Wiley-Liss, Inc.     

Cryptochrome 1b: a possible inducer of visual lateralization in pigeons?

The visual system of adult pigeons shows a lateralization of object discrimination with a left hemispheric dominance on the behavioural, physiological and anatomical levels. The crucial trigger for the establishment of this asymmetry is the position of the embryo inside the egg, which exposes the right eye to light falling through the egg shell. As a result, the right‐sided retina is more strongly stimulated with light during embryonic development. However, it is unknown how this embryonic light stimulation is transduced to the brain as rods and cones are not yet functional. A possible solution could be the blue‐light‐sensitive molecule cryptochrome 1 (Cry1), which is expressed in retinal ganglion cells (RGCs) of several mammalian and avian species. RGCs have been shown to be functional during the time of induction of asymmetry and possess projections to primary visual areas. Therefore, Cry1‐containing RGCs could be responsible for induction of asymmetry. The aim of this study was to identify the expression pattern of the Cry1 subtype Cry1b in the retina of embryonic, post‐hatch and adult pigeons by immunohistochemical staining and to show whether Cry1b‐containing RGCs project to the optic tectum. Cry1b‐positive cells were indeed mainly found in the RGC layer and to lesser extent in the inner nuclear layer at all ages, including the embryonic stage. Tracing in adult animals revealed that at least a subset of Cry1b‐containing RGCs project to the optic tectum. Thus, Cry1b‐containing RGCs within the embryonic retina could be involved in the induction of asymmetries in the visual system of pigeons.     

Abnormal ipsilateral visual field representation in areas 17 and 18 of hypopigmented cats

We compared the central projections of retinal ganglion cells in temporal retina and the cortical representation of visual fields in areas 17 and 18 in cats with various hypopigmentation phenotypes (albino, heterozygous albino, Siamese, and heterozygous Siamese). In all cats studied, we found that the extent of abnormal ipsilateral visual field representation varied widely, and more of the ipsilateral visual field was represented in area 18 than in area 17. The greatest degree of ipsilateral visual field representation was found in albino cats, followed by Siamese, heterozygous albino and heterozygote Siamese cats, respectively. Additionally, in the different groups there was wide variation in the numbers of contralaterally projecting alpha and beta ganglion cells in temporal retina. In all cases, however, contralaterally projecting alpha cells were found to extend further into temporal retina than beta cells. We found that in each cat studied, the maximum extent of the abnormal ipsilateral visual field representation in areas 18 and 17 corresponded to the location of the 50% decussation line (i. e., the point where 50% of the ganglion cells in temporal retina project to the contralateral hemisphere) for alpha and beta cells, respectively, for that cat. Our results suggest that the extent of the abnormal visual field representations in visual cortex of hypopigmented cats reflects the extent of contralaterally projecting retinal ganglion cells in temporal retina. © 1995 Wiley-Liss, Inc.     

Different regional specializations of neurons in the ganglion cell layer and inner plexiform layer of the California horned shark, Heterodontus francisci

We have described a population of neurons in the retina of a shark, Heterodontus francisci, which is precisely aligned within the inner plexiform layer (IPL) and which differs from neurons in the ganglion cell layer (GCL) in soma size and topographical distribution. GCL neurons are relatively small and form a horizontally oriented visual streak; IPL neurons are significantly larger and form a circular specialization in the far temporal retina. Thus, it appears that there are two distinct retinal specializations in Heterodontus: one subserving frontal vision and one which provides a panoramic view of the lateral visual field.     

Cell transplantation to replace retinal ganglion cells faces challenges – the Switchboard Dilemma

The mammalian retina displays incomplete intrinsic regenerative capacities; therefore, retina degeneration is a major cause of irreversible blindness such as glaucoma, age-related macular degeneration and diabetic retinopathy. These diseases lead to the loss of retinal cells and serious vision loss in the late stage. Stem cell transplantation is a great promising novel treatment for these incurable retinal degenerative diseases and represents an exciting area of regenerative neurotherapy. Several suitable stem cell sources for transplantation including human embryonic stem cells, induced pluripotent stem cells and adult stem cells have been identified as promising target populations. However, the retina is an elegant neuronal complex composed of various types of cells with different functions. The replacement of these different types of cells by transplantation should be addressed separately. So far, retinal pigment epithelium transplantation has achieved the most advanced stage of clinical trials, while transplantation of retinal neurons such as retinal ganglion cells and photoreceptors has been mostly studied in pre-clinical animal models. In this review, we opine on the key problems that need to be addressed before stem cells transplantation, especially for replacing injured retinal ganglion cells, may be used practically for treatment. A key problem we have called the Switchboard Dilemma is a major block to have functional retinal ganglion cell replacement. We use the public switchboard telephone network as an example to illustrate different difficulties for replacing damaged components in the retina that allow for visual signaling. Retinal ganglion cell transplantation is confronted by significant hurdles, because retinal ganglion cells receive signals from different interneurons, integrate and send signals to the correct targets of the visual system, which functions similar to the switchboard in a telephone network – therefore the Switchboard Dilemma.     

The centrifugal visual system of vertebrates: a comparative analysis of its functional anatomical organization

The present review is a detailed survey of our present knowledge of the centrifugal visual system (CVS) of vertebrates. Over the last 20 years, the use of experimental hodological and immunocytochemical techniques has led to a considerable augmentation of this knowledge. Contrary to long-held belief, the CVS is not a unique property of birds but a constant component of the central nervous system which appears to exist in all vertebrate groups. However, it does not form a single homogeneous entity but shows a high degree of variation from one group to the next. Thus, depending on the group in question, the somata of retinopetal neurons can be located in the septo-preoptic terminal nerve complex, the ventral or dorsal thalamus, the pretectum, the optic tectum, the mesencephalic tegmentum, the dorsal isthmus, the raphé, or other rhombencephalic areas. The centrifugal visual fibers are unmyelinated or myelinated, and their number varies by a factor of 1000 (10 or fewer in man, 10,000 or more in the chicken). They generally form divergent terminals in the retina and rarely convergent ones. Their retinal targets also vary, being primarily amacrine cells with various morphological and neurochemical properties, occasionally interplexiform cells and displaced retinal ganglion cells, and more rarely orthotopic ganglion cells and bipolar cells. The neurochemical signature of the centrifugal visual neurons also varies both between and within groups: thus, several neuroactive substances used by these neurons have been identified; GABA, glutamate, aspartate, acetylcholine, serotonin, dopamine, histamine, nitric oxide, GnRH, FMRF-amide-like peptides, Substance P, NPY and met-enkephalin. In some cases, the retinopetal neurons form part of a feedback loop, relaying information from a primary visual center back to the retina, while in other, cases they do not. The evolutionary significance of this variation remains to be elucidated, and, while many attempts have been made to explain the functional role of the CVS, opinions vary as to the manner in which retinal activity is modified by this system.