Interactions between tectal radial cells in the red-eared turtle, Pseudemys scripta elegans: an analysis of tectal modules.

The optic tectum is a major subdivision of the visual system in reptiles. Previous studies have characterized the laminar pattern, the neuronal populations, and the afferent and efferent connections of the optic tectum in a variety of reptiles. However, little is known about the interactions that occur between neurons within the tectum. This study describes two kinds of interactions that occur between one major class of neurons, the radial cells, in the optic tectum of Pseudemys using Nissl, Golgi and electron microscopic preparations. Radial cells have somata which bear long, radially oriented apical dendrites from their upper poles and short, basal dendrites from their lower poles. They are divided into two populations on the basis of the distribution of their somata in the tectum. Deep radial cells have somata densely packed in the stratum griseum periventriculare. Their plasma membranes form casual appositions. Middle radial cells have somata scattered throughout the stratum griseum centrale and stratum fibrosum et griseum superficiale and do not contact each other. The apical dendrites of both populations of radial cells participate in vertically oriented, dendritic bundles. The plasma membranes of the dendrites in these bundles form casual appositions in the deeper tectal layers and chemical, dendrodenritic synapses within the stratum fibrosum et griseum superficiale. The synapses have clear, round synaptic vesicles and slightly asymmetric membrane densities. Thus, radial cells interact via both casual appositions and chemical synapses. These interactions suggest that radial cells may form a basic framework in the tectum. Because both populations of radial cells extend into the stratum fibrosum et griseum superficiale and stratum opticum, they may receive input from some of the same tectal afferent systems. Because the deep radial cells alone have somata and dendrites in the deep tectal layers, they may receive additional inputs that the middle radial cells do not. Neurons in the two populations interact via chemical dendrodentritic synapses, thereby forming vertically oriented modules in the tectum.     

Glutamate-like immunoreactivity in chick cerebellum and optic tectum

Glutamate was coupled via glutaraldehyde to bovine serum albumin. The conjugate was used for raising specific anti-glutamate antibodies. The purified antibody was used for immunostaining of chick cerebellum and optic tectum. Staining was intense in the molecular layer and in cell bodies of the granule cell layer. In the optic tectum a diffuse staining was detected in the superficial layers of stratum griseum fibrosum superficiale and in cell bodies especially in the layers a and e. Large cell bodies located in the stratum griseum centrale were also stained.     

墨龙与红鲫的视网膜和视盖解剖结构比较

墨龙是一种由红鲫进化来的龙睛种金鱼(Carassius auratus)。随机取体长10—12 cm, 重约35 g的墨龙和红鲫各4尾, 解剖取出整个眼球及脑, 并常规石蜡切片, HE染色。在光学显微镜下观察墨龙和红鲫的视网膜、视盖系统的显微结构变化并比较各层厚度, 发现与红鲫相比, 墨龙视网膜的总厚度下降29.9%, 其中外网状层厚度增加2.5%、内网状层厚度增加11.8%; 而内核层厚度下降21.6%、外核层厚度降低35.6%, 神经节细胞层、杆锥层也变薄, 且后两者分层不规则; 墨龙视盖壁整体厚度增加28.9%, 其中除围脑室层厚度减少22.6%外, 中央纤维层厚度增加12.8%, 中央细胞层厚度增加30.6%, 表面纤维层厚度增加21.9%, 且纤维远较红鲫密集, 视神经层厚度增加91.7%, 边缘层厚度增加35.6%。结果表明长期的人工选择不但改变了墨龙的外形, 而且使其中枢神经组织结构也发生了较大变化, 并推测墨龙的眼球直径及视网膜面积较大, 从而导致自视网膜传入视盖的纤维增多, 是视网膜和视盖中的传递神经冲动的神经元、神经纤维所在层段增厚的主要原因; 同时墨龙视网膜中色素上皮层向杆锥层交错对插, 富含神经元的视网膜外核层、内核层以及视盖中的围脑室层厚度也降低, 可以减少因视网膜面积大而造成的强光伤害; 此外由于墨龙的围脑室层厚度降低, 导致其游动及平衡能力较红鲫差。     

Laminar distribution of current sources in the rabbit superior colliculus during optic nerve stimulation

The electric field of evoked potentials developing in the rabbit superior colliculus in response to single electrical stimulation of the optic nerve was investigated. Regions of localization of sources and sinks of synaptic current creating the evoked potential were discovered by the current source density analysis method. Sinks with the shortest latency appeared in the depth of str. griseum superficiale, after which activity created by two dipoles appeared: The sink of one dipole was located in the upper part of str. griseum superficiale, the sink of the other in the lower part. The sinks thus found corresponded in their temporal characteristics and location to the principal components of the combined evoked potential. On the basis of these data the order of activation of the various systems of fibers and synapses responsible for evoked potential formation can be traced.     

Regional effect of monosodium-L-glutamate on the superficial layers of superior colliculus in rat

Summary Systemic administration of monosodium-1-gluta-mate by single injections of 4 mg/g body weight in infant rats (2–10 days of age) results in acute swelling of cytoplasm and nuclear pyknosis of neurons in the stratum zonale and stratum griseum superficiale of the superior colliculus. Multiple daily doses of 4 mg/g body weight monosodium-1-glutamate result in an almost complete loss of neurons in these two superficial layers. The deeper layers appear not to be affected. No pathological effects were observed in the lateral geniculate body or pretectal complex.Light-and electron-microscopic studies reveal that the optic nerves are remarkably shrunken and many myelinated as well as unmyelinated axons are lost. Injection of ³Hproline into the vitreous body of one eye results in limited transport to the suprachiasmatic nucleus, lateral geniculate body and to lateral portions of the superior colliculus.The small percentage of intact axons in the optic nerve, as well as the limited proline transport from the eye, suggest that administration of monosodium-1-glutamate leaves intact some optic fibers, a portion of which belongs to the retinohypothalamic tract.     

Distribution of calbindinlike immunoreactive structures in the optic tectum of normal and eye-enucleated cyprinid fish

Summary Using the ABC immunohistochemical method, we investigated the distribution of calbindinlike immunoreactive structures in the optic tectum of normal fish, Tinca tinca, and from normal and unilaterally eye-enucleated fish, Cyprinus carpio. In nonoperated individuals of both species the optic tectum contained numerous immunoreactive neurons with strongly positive somata located in the stratum periventriculare and a thick immunolabeled dendritic shaft ascending radially toward the stratum fibrosum et griseum superficiale. The retinorecipient layers contained many fibrous immunoreactive structures. Some varicose fibers, isolated or in small bundles, were localized to the stratum album centrale, especially in the dorsal tectal half. Unilateral eye removal produced the disappearance of the immunoreactive fibrous structures located in the retinorecipient layers of the tectum contralateral to the enucleation. The present work shows that calbindinlike immunoreactive substances are localized in specific neural circuits of the fish optic tectum and suggests that the calbindin-like immunoreactive fibers in the retinorecipient strata are of retinal origin.     

Neuronal and synaptic organization in the gravity receptor system of the statocyst of Octopus vulgaris

Summary The optic tectum of Calamoichthys calabaricus (Polypteriformes) shows a relatively complex vertical stratification, with six main layers and a varied neuronal typology. In particular, pyriform neurons in the well developed stratum griseum periventriculare and some multipolar neurons in the stratum griseum profundum represent the efferent elements of the tectum, while the optic and lemniscal inputs to the tectum converge in the plexiform sublayers of the stratum fibrosum et griseum superficiale. In the circuitry of the tectum some modulation is achieved by some of the polymorphic cells of the stratum griseum internum and by the horizontal cells of the outer layers. Notwithstanding some differences with respect to the teleost optic lobe (i.e., the absence of a torus longitudinalis; the lack of a stratum fibrosum marginale; the modest size of the stratum fibrosum profundum; the paucity of neurons in the stratum fibrosum et griseum superficiale; and the ill-defined separation of the layers of the afferent and efferent fibers), the optic tectum of Calamoichthys resembles the mesotectal type characteristic of teleosts, anurans and reptiles. It exhibits higher degree of organization than the optic tectum of the Chondrostei.     

Retinal projections in the catfish,Mystus vittatus (Bloch) as revealed by tracer studies with horseradish peroxidase

Summary The retinal efferents of the catfish, Mystus vittatus, were investigated with the use of the horseradish peroxidase (HRP) technique. Most retinal fibres extended contralateral to the eye that had received HRP label, while a few fascicles projected to the ipsilateral side without decussation in the optic chiasma. The contralateral fibres projected to the suprachiasmatic nucleus, the nucleus opticus dorsolateralis, the nucleus of the posterior commissure, the nucleus geniculatus lateralis, pretectal nuclear complex, and to two layers of the optic tectum, i.e., stratum fibrosum et griseum superficiale and stratum griseum centrale. The accessory optic tract arose from the inner area of the optic tract and extended ventromedially to the accessory optic nucleus. The ipsilateral fascicles projected to almost all the above mentioned nuclei, but these projections were comparatively sparse. The ipsilateral retinal projection was restricted to the rostral tectum.     

Distribution of MAP1A,MAP1B,and MAP2A&;B during layer formation in the optic tectum of developing chick embryos

The expression patterns of three microtubule-associated proteins (MAP1A, MAP1B, and MAP2A&B) were investigated in the developing optic tectum. Expression of MAP1B and middle-molecular-weight peptide of neurofilament (NF-M) was first observed in the same mesencephalic cells on day 3 of incubation, indicating that neuroblasts had been produced. At day 5, MAP1A and MAP2A&B expression appeared in the cellular layer containing the first neuroblasts that differentiate into large multipolar cells. The NF-M⁺ neurites in the striatum album centrale (SAC) and the striatum opticum (SO) were MAP1B⁺ up to day 19, but the intensity of MAP1B immunoreactivity decreased with development. All three MAPs were expressed in large multipolar neurons in the developing stratum griseum centrale from the beginning of maturation. Stratum griseum et fibrosum centrale cellular layers, containing radially arranged piriform neurons, were MAP1A^–/MAP2A&B^– on day 11 but became MAP1A⁺/MAP2A&B⁺ during later stages. These results suggest that the timing of MAP expression in neuronal maturation of large multipolar cells differs from that of piriform cells. The expression of MAPs has revealed specific cellular events in the developing optic tectum. Based on our observations, the development of the optic tectum can be divided into four periods.     

Presynaptic dendrite cells and two other classes of neurons in the superficial layers of the superior colliculus of the chimpanzee

In the superior colliculus of chimpanzee, three classes of neurons can be identified by ultrastructural criteria. They are 1) marginal cells located in the stratum zonale, 2) collicular relay cells in the stratum griseum superficiale and 3) presynaptic dendrite (PSD) cells, i.e., neurons with presynaptic specializations in soma and/or dendrites. PSD cells are the smallest neurons in the stratum griseum superficiale; they have a relatively large, deeply infolded nucleus and a small rim of cytoplasm rich in free ribosomes. PSD cells are sufficiently different from the two other classes of neurons to be reliably identified at the ultrastructural level. They closely resemble presynaptic neurons as described in the lateral geniculate nucleus of other mammalian species. Presynaptic dendrites in continuity with PSD cells are rich in organelles, especially ribosomal cluster, and establish en passage contact with other dendrites. Another type of presynaptic dendrite, poor in organelles, except for bundles of microtubules, could not be traced back to its parent neuron. Homo- or heterogeneity of PSD cells is discussed. No amxon was traced from a PSD cell.     

Ultrastructural changes in the superficial layers of the superior colliculus in Galago crassicaudatus (primates) after eye enucleation

Summary Several types of terminals were found in the three superficial collicular layers of Galago. At least two axon terminals with round vesicles (R1 and R2) could be distinguished on the basis of vesicle packing and electron density of the cytoplasmic and mitochondrial matrices. R1 axon terminals were characterized by aggregations of vesicles in an electron lucent cytoplasm and mitochondria with a relatively dark matrix, while in R2 axon terminals the vesicles were more evenly distributed in an electron dense cytoplasm and the mitochondrial matrix was pale. R2 endings occurred in clusters in the stratum griseum superficiale; they were absent in the stratum zonale. R1 endings were found in all three superficial collicular layers. Both types of R terminals made asymmetrical contacts with small dendrites, dendritic spines and F profiles. Profiles containing flattened vesicles and establishing symmetrical contacts were numerous, and many could be identified as dendrites by accepting as criteria for dendrites evenly spaced microtubules, clusters of ribosomes and the fact that these F profiles were postsynaptic to other terminals. F terminals were presynaptic to other F profiles, dendrites and somata; they were postsynaptic to R terminals and took part in serial synapses. Dendrodendritic contacts were frequent, somatodendritic contacts rare. After eye enucleation most R2 axon terminals underwent the electron dense degenerative reaction. The degeneration process was a lengthy one; many degenerating boutons were found 30 days after axotomy and some persisted up to 180 days postoperatively. There was strong indication that the superior colliculus received more crossed than uncrossed retinofugal fibers. The crossed and uncrossed retinocollicular axons terminated in two different substrata of the stratum griseum superficiale.This study was supported by N.I.H. Grant RR-00165 to Yerkes Regional Primate Research Center and N.I.H Grant EY 00638-03 to J. Tigges. — The opportunity to use the electron microscopic facilities of the Fernbank Science Center for the initial stage of this work is gratefully acknowledged.     

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.     

Neurons with complex receptive fields in the stratum griseum centrale of the zebra finch (Taeniopygia guttata castanotis Gould) optic tectum

The electrophysiological and morphological features of visually driven neurons of the stratum griseum centrale of the zebra finch optic tectum were studied by extracellular recording and staining techniques. Stratum griseum centrale neuron responses are sustained in most cases. Receptive fields are big, up to 150 degrees of the visual field. The excitatory center (hot spot) varies in size from 1 degrees to 15 degrees. It can be mapped by small static stimuli, adapts slower than the surround, and has a shape comparable to the excitatory fields of upper-layer neurons. In contrast, the big surround shows responses only to small moving objects which elicit a typical pattern of alternating bursts and silent periods. Alternatively, the same stimuli elicit long-lasting bursts followed by strong adaption. Anatomically, stratum griseum centrale neurons are characterized by far reaching dendrites which terminate with "bottlebrush"-like endings in the upper retinorecipient layers. In addition, they are connected with retinorecipient structures by an interneuron located between layers 10 and 11. The role of the structure of inputs for the organization of the receptive fields is discussed.     

Retinal projections in the chain pickerel (Esox niger Lesueur)

Summary The retinal projections inEsox niger, as determined with the aid of a modified cobalt-lysine method, are considerably more extensive in the diencephalon and pretectum than in other teleost fishes so far examined. Although most retinal axons terminate contralaterally, rare fibers can be traced to the same aggregates ipsilaterally. The retinohypothalamic projection appears larger than hitherto reported in teleosts, and the dorsomedial optic tract issues fibers to a series of cell clusters extending from the rostral thalamus to mid-torus levels. A retinal projection to a presumed ventrolateral optic nucleus (VLO) is described for the first time in a teleost. Other targets of retinal fibers include the nucleus geniculatus lateralis ipse of Meader (GLI), the pretectal nucleus (P), the cortical nucleus and a well-developed ventromedial optic nucleus (VMO). The projection to the optic tectum is principally to the stratum fibrosum et griseum superficiale (SFGS) and stratum marginale (SM), but a considerable number of axons also course through the stratum album centrale (SAC) before terminating there or piercing the stratum griseum centrale (SGC) and terminating in SFGS. Rare terminal arborizations of retinal fibers were also observed in stratum griseum centrale (SGS) and in the stratum griseum periventriculare (SGC) in restricted portions of the tectum. Because of the relatively large size of the visual structures inE. niger it is a potentially useful model for future experimental studies on the visual system.     

Excitation and inhibition in the visual center of Rana temporaria

Analysis of postsynaptic unit responses in the visual center ofRana temporaria showed that optic nerve fibers with high and low conduction velocities usually converge on a single neuron of the tectum opticum (TO). In response to stimulation of the optic nerve a complex depolarization potential consisting of 3 (or possibly 4) EPSPs was recorded in one group of neurons; these EPSPs were probably generated through excitation of several groups of afferent fibers. Either an increase or a decrease in the EPSPs can be observed in the TO neurons in response to repetitive and paired stimulation of the optic nerve. Postsynaptic inhibitory responses of some TO neurons, probably of direct and recurrent origin, are discussed.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 3, No. 6, pp. 637–643, November–December, 1971.     

Neurophysiological properties of magnetic cells in the pigeon's visual system

Summary Single unit electrical activity was recorded extracellularly in the nucleus of the basal optic root (nBOR) and in the optic tectum under earth-strength magnetic stimulation. Units in the nBOR which were stimulated while the eyes were illuminated by light of different wavelengths exhibited peaks of magnetic responsiveness at 503 nm and 582 nm.Magnetically directional selective cells were found in the stratum griseum et fibrosum superficiale of the optic tectum. They also showed directional selectivity to dynamic photic stimuli. Response peaks varied with the orientation of the pigeon in the horizontal plane. This confirmed that the magnetic responses contained directional information. The results suggest that the receptor and neural organisation of the pigeon's visual system provides an adequate substrate for the detection and elaboration of magnetic compass information.     

Immunoreactive endozepine-like peptides in the brain and pituitary of the Atlantic hagfish, Myxine glutinosa

Endozepines are a family of peptides capable of displacing benzodiazepines from their specific binding sites, to which belong the diazepam-binding inhibitor and the octadecaneuropeptide (ODN). This paper reports the distribution of ODN-related peptides, investigated for the first time by immunocytochemistry, in different brain and pituitary regions of the Atlantic hagfish, Myxine glutinosa. Immunoreactive ODN-like material was found in the telencephalon at the level of bundles of different olfactory nerve fibres. Moreover, at the level of the pallium, immunoreactive multipolar neurons were observed in the pars parvocellularis of the stratum griseum superficialis. Similar immunopositive nerve cell bodies were found in the nucleus medialis of the central prosencephalic complex. In the mesencephalon, few immunoreactive neurons lining and contacting the mesencephalic ventricle were detected; such nerve cells could be involved in the regulation of cerebrospinal fluid homeostasis. Dorsally in the mesencephalon, numerous ODN-containing cell bodies were present in the area praetectalis. The rhomboencephalon was immunostained only in the octavolateral area and in the nucleus motorius magnocellularis of the trigeminal nerve. Furthermore, ODN immunoreactivity was also present in the nerve cells of ganglia of the ophthalmic division of the trigeminal nerve complex. The immunocytochemical patterns described here in the brain of M. glutinosa suggest an involvement of ODN-like peptides as neuromodulators in sensory pathways, such as olfactory and visual. Finally, ODN-like substances were localized in discrete populations of adenohypophysial cells and in tanycytes lining the neurohypophyseal walls, suggesting for endozepines a paracrine and/or endocrine control of pituitary hormones release and a neurohormone role respectively. These results could give new insights into the chemioarchitecture of the brain of myxinoids.     

Molecular mechanisms separating two axonal pathways during embryonic development of the avian optic tectum

During embryonic development of the avian optic tectum, retinal and tectobulbar axons form an orthogonal array of nerve processes. Growing axons of both tracts are transiently very closely apposed to each other. Despite this spatial proximity, axons from the two pathways do not intermix, but instead restrict their growth to defined areas, thus forming two separate plexiform layers, the stratum opticum and the stratum album centrale. In this study we present experimental evidence indicating that the following three mechanisms might play a role in segregating both axonal populations: Retinal and tectobulbar axons differ in their ability to use the extracellular matrix protein laminin as a substrate for axonal elongation; the environment in the optic tectum is generally permissive for retinal axons, but is specifically nonpermissive for tectobulbar axons, resulting in a strong fasciculation of the latter; and growth cones of temporal retinal axons are reversibly inhibited in their motility by direct contact with the tectobulbar axon's membrane.     

Pax genes in development and maturation of the vertebrate visual system: implications for optic nerve regeneration

Pax genes play a pivotal role in development of the vertebrate visual system. Pax6 is the master control gene for eye development: ectopic expression of Pax6 in Xenopus laevis and Drosphila melanogaster leads to the formation of differentiated eyes on the legs or wings. Pax6 is involved in formation of ganglion cells of the retina, as well as cells of the lens, iris and cornea. In addition Pax6 may play a role in axon guidance in the visual system. Pax2 regulates differentiation of the optic disk through which retinal ganglion cell axons exit the eye. Furthermore, Pax2 plays a critical role in development of the optic chiasm and in the guidance of axons along the contralateral or ipsilateral tracts of the optic nerve to visual targets in the brain. During development Pax7 is expressed in neuronal cells of one of the major visual targets in the brain, the optic tectum/superior colliculus. Neurons expressing Pax7 migrate towards the pia and concentrate in the stratum griseum superficiale (SGFS), the target site for retinal axons. Together, expression of Pax2, 6 and 7 may guide axons during formation of functional retinotectal/collicular projections. Highly regulated Pax gene expression is also observed in mature animals. Moreover, evidence suggests that Pax genes are important for regeneration of the visual system. We are currently investigating Pax gene expression in species that display a range of outcomes of optic nerve regeneration. We predict that such information will provide valuable insights for the induction of successful regeneration of the optic nerve and of other regions of the central nervous system in mammals including man.