Element Distribution in Visual System,the Optic Chiasma,Lateral Geniculate Body,and Superior Colliculus

To elucidate compositional changes of the visual system with aging, the authors investigated age-related changes of elements in the optic chiasma, lateral geniculate body, and superior colliculus, relationships among their elements, relationships among their brain regions from a viewpoint of elements, and gender differences in their elements by direct chemical analysis. After ordinary dissection at Nara Medical University was finished, the optic chiasmas, lateral geniculate bodies, and superior colliculi were resected from identical cerebra of the subjects. The subjects consisted of 14 men and 10 women, ranging in age from 75 to 96 years (average age = 85.6 ± 5.9 years). After ashing with nitric acid and perchloric acid, element contents were determined by inductively coupled plasma-atomic emission spectrometry. As the result, the average content of P was significantly higher in the optic chiasma and superior colliculus compared with the lateral geniculate body. Regarding age-related changes of elements, no significant changes with aging were found in seven elements of the optic chiasma, lateral geniculate body, and superior colliculus in the subjects more than 75 years of age. The findings that with regard to the relationships among elements, there were extremely significant direct correlations between Ca and Zn contents and significant inverse correlations between Mg and Na contents were obtained in common in all of the optic chiasma, lateral geniculate body, and superior colliculus. It was examined whether there were significant correlations among the optic chiasma, lateral geniculate body, and superior colliculus in the seven elements and the following results were obtained: There were significant direct correlations between the optic chiasma and lateral geniculate body in both the P and Mg contents; there was a significant direct correlation between the optic chiasma and superior colliculus in the Fe content; and a significant direct correlation was found between the lateral geniculate body and superior colliculus in the Mg content. Regarding the gender differences in elements, it was found that both the Ca and Zn contents of the lateral geniculate body were significantly higher in women than in men.     

Development of muscarinic cholinergic receptor binding in the visual system of monocularly deprived and dark reared rats

In 25 day old rats monocularly deprived by unilateral eyelid suture on postnatal day 10 (MD), [³H]quinuclidinyl benzylate (³H-QNB) binding was significantly reduced in the visual cortex (VC) of both sides, but elevated in both superior colliculi (SC). Muscarinic receptor binding in the frontal cortex (FC), a non-visual brain area, in the lateral geniculate nucleus (LGN), and in the retina was not affected. In 25 day old rats raised in complete darkness from birth (DR) similar changes in³H-QNB binding were found in VC and SC. However, binding levels were also decreased in the FC and significantly increased in the retina. In adult (6 month old) MD and DR rats the differences in³H-QNB binding as compared to age-matched controls had disappeared completely in all visual brain areas studied. Detailed Scatchard analyses indicate that the alterations in the³H-QNB binding were due to changes in receptor number only.This paper is dedicated to Dr. Derek Richter on his seventy-fifth birthday.     

Formation of slow background activity in different parts of the visual analyzer following section of half the tectum mesencephali

Significant changes in the formation of electrical activity rhythms have been revealed in the lateral geniculate body, superior colliculus and visual cortex during section of one half of midbrain operculum in cats anesthetized with nembutal. It was determined that all changes in slow activity generation in the lateral geniculate body, superior colliculus are reflected in changes in the formation of electrical activity of the visual cortex. It is suggested that lateral geniculate body and superior colliculus may be involved in the generation of some electrical activity rhythms of the visual cortex.     

The Effect of Visual Deprivation on β-Adrenergic Receptors in the Visual Centres of the Rat Brain

The levels of binding of [3H]dihydroalprenolol to beta-adrenergic receptors in the visual centres and frontal cortex from brains of control, dark-reared and monocularly deprived rats were compared. Receptor binding is changed in monocularly deprived rats in the lateral geniculate nuclei and superior colliculi of both sides. Scatchard analyses indicated that the changes in the [3H]dihydroalprenolol binding in the lateral geniculate nuclei were due to alterations in both receptor affinity and receptor number. No effect of dark-rearing could be detected.     

Effect of Monocular Deprivation on Uptake and Binding of [3H]Glutamate in the Visual System of Rat Brain

Abstract: [³H]Glutamate uptake and binding studies were performed in the visual cortices, lateral geniculate nuclei (LGN), and superior colliculi of 3-month-old rats with one eyelid surgically closed from postnatal day 10 (monocular deprivation). Uptake and binding were highest in the lateral geniculate nucleus followed by the visual cortex (69% and 15%, respectively compared to LGN values) and the superior colliculus (32% and 59% of LGN values). Monocular deprivation did not affect [³H]glutamate uptake in any of the visual regions examined. However, a 46% decrease in [³H]glutamate binding in the lateral geniculate nucleus ipsilateral to the sutured eye was detected. Binding levels in other regions were not affected.     

Intra-analyzer temporary connections in the visual system]

The possibility of setting up intraanalyser temporary connections integrating the activity of various elements within the central part of the visual analyser at a neuronal level was studied in acute experiments on unanaesthetized immobilized cats. In the given model of temporary connection the unit activity in the lateral geniculate bodies (LGB) was investigated. Electrical stimulation of the superior colliculi was used as a conditioned stimulus, and illumination of the receptive fields of the studied LGB neurones, as an unconditioned one. In the process of conditioning, 10 to 11% of learning elements were revealed in LGB. The possibility is discussed of integration of learning elements into a definite microsystem achieving the process of temporary connection closing in the visual analyser in the course of conditioning.     

Corticalization of two divisions of the visual system during vertebrate evolution

Data on the evolution of the visual system in vertebrate phylogeny are described. Visual projections are demonstrated in the telencephalon of cyclostomata (lampreys). The existence of a retino-thalamo-telencephalic pathway is demonstrated in elasmobranchs (skates). Two visual pathways are present in amphibians (frogs) and reptiles (turtles): retino-thalamo-telencephalic and retino-tecto-thalamo-telencephalic, and these overlap partly at the thalamic level in the lateral geniculate nucleus and completely in the telencephalon. In turtles the earliest visual and tectal impulses relay on their way to the telencephalon in the lateral geniculate body, and later impulses relay in the nucleus rotundus. In mammals (rats) visual tecto-cortical connections are seen; judging from the latent period of potentials arising in the visual cortex in response to stimulation of the superior colliculi these connections have one synaptic relay in the thalamus. The much shorter latent periods of visual evoked potentials recorded in the tectum of the monkey than in turtles (under identical chronic experimental conditions) confirm the views of morphologists on the progressive development of the tectal division of the visual system in vertebrate phylogeny. It is concluded that corticalization of both divisions of the visual system, i.e., the existence of telencephalic representation, appears in the early stages of vertebrate evolution.     

Retinal projections of the pigmented guinea pig

Using 3H-proline autoradiography we found in the pigmented guinea pig that retinal fibres terminated bilaterally in the suprachiasmatic, lateral geniculate, some pretectal nuclei and in the superior colliculi. The medial, lateral and dorsal terminal nuclei of the accessory optic system appeared to receive only contralateral retinal fibres. The retinal projections of the guinea pig follow the general plan recognized in the visual system of rodents demonstrated by modern tract-tracing techniques.     

Development of GABAA receptors in the central visual structures of rat brain. Effect of visual pattern deprivation

The postnatal development of high-affinity 3H-muscimol binding to GABAA receptors was studied in the lateral geniculate nucleus, superior colliculus, frontal and visual cortex of the rat brain. In the lateral geniculate nucleus 3H-muscimol binding rises from day 10 through day 37 reaching the highest value during the entire development followed by a slight decrease until adulthood. In the superior colliculus 3H-muscimol binding increased continuously from day 10 through day 37, and then decreased until day 50 reaching the adult value. In the visual and frontal cortex, binding reached the highest levels on days 14 and 25, respectively, persisted until day 37 followed by a slight decrease until adulthood. The ontogeny of 3H-muscimol binding sites in the visual regions does not essentially differ from that in other brain regions, suggesting that the appearance of 3H-muscimol binding sites in the visual system is not correlated with the functional maturation of the visual system. Unilateral eyelid closure from day 11 until day 25 did not affect the development of GABAA receptors in any of the central visual regions examined, indicating the lack of environmentally controlled mechanism.     

Cholesterol is a component of the rapid phase of axonal transport

Twenty-two-day-old rats were injected intraocularly with [3H]acetate and killed between 1 hr and 35 days later. Cholesterol was isolated from the retinas, optic tracts, lateral geniculate bodies, and superior colliculi. Within the retina, radioactivity was rapidly incorporated into cholesterol with maximal labeling present one hour after injection. Transported labeled cholesterol (contralaterally corrected for systemic background labeling) was present in the superior colliculus by three hours. Radioactive cholesterol accumulated in all visual structures throughout the 35-day period, but the rate of accumulation was maximal at about the time of arrival of the initial pulse of radioactivity. Colchicine treatment of the retina blocked transport of cholesterol but not its synthesis by the retina. The results indicate that cholesterol is rapidly transported in the visual system and also released from the retina for a prolonged period after its synthesis.     

Amplitude-temporal characteristics of evoked responses of the visual analyzer to stimuli of growing intensity

A study was made on cats of the dependence of latency, peak latency, amplitudes and slopes of I and II phases of primary evoked potentials in the chiasm, the colliculi, the lateral geniculate body and visual cortex on the intensity of the photic stimulus in the range of intensities of II orders above the threshold. Practically in the whole examined range, the logarithmic connection is retained, testifying to the extremely wide possibility of the visual system to discriminate a signal in securing a reflex act.     

Influence of early visual deprivation on regional activity of brain ATPases in developing rats

The effects of light deprivation from birth to 40 days of age on the development of Na-K-ATPase and Mg-ATPase activity, enzymes significantly involved in cerebral energy exchange, ion transport, and synaptic function, were investigated in visual and non-visual brain areas of the rat. Although both enzymes generally showed a progressive depression with age in light-deprived rats, Na-K-ATPase was more depressed than Mg-ATPase, and significant effects were confined to the superior colliculi, visual cortex, frontal cortex and hypothalamus. A disparate developmental pattern was evidenced in Na-K-ATPase activity in the visual cortex, where it was higher than control values at day 10 but lower by day 40, and in the hypothalamus, where it was lower on days 10 and 25 but significantly higher on day 40. The depression of Mg-ATPase in the hypothalamus of light-deprived rats at all ages and the activation of Na-K-ATPase in this structure is interpreted to mean that discrete alterations may have occurred in neurosecretory functions of the hypothalamus, known to be responsive to light. Transferring dark-reared animals to normal light-cycle conditions at day 25 affected only Mg-ATPase in the visual cortex and Na-K-ATPase in the hypothalamus, both enzymes showing a significant increase by day 40 over values in continually light-deprived animals. These findings confirm that early light deprivation is associated with important biochemical and neuroendocrine changes that persist into adulthood.     

Axonal transport of the mitochondria-specific lipid, diphosphatidylglycerol, in the rat visual system

Rats 24 d old were injected intraocularly with [2-3H]glycerol and [35S]methionine and killed 1 h-60 d later. 35S label in protein and 3H label in total phospholipid and a mitochondria-specific lipid, diphosphatidylglycerol(DPG), were determined in optic pathway structures (retinas, optic nerves, optic tracts, lateral geniculate bodies, and superior colliculi). Incorporation of label into retinal protein and phospholipid was nearly maximal 1 h postinjection, after which the label appeared in successive optic pathway structures. Based on the time difference between the arrival of label in the optic tract and superior colliculus, it was calculated that protein and phospholipid were transported at a rate of about 400 mm/d, and DPG at about half this rate. Transported labeled phospholipid and DPG, which initially comprised 3-5% of the lipid label, continued to accumulate in the visual structures for 6-8 d postinjection. The distribution of transported material among the optic pathway structures as a function of time differed markedly for different labeled macromolecules. Rapidly transported proteins distributed preferentially to the nerve endings (superior colliculus and lateral geniculate). Total phospholipid quickly established a pattern of comparable labeling of axon (optic nerve and tract) and nerve endings. In contrast, the distribution of transported labeled DPG gradually shifted toward the nerve ending and stabilized by 2-4 d. A model is proposed in which apparent "transport" of mitochondria is actually the result of random bidirectional saltatory movements of individual mitochondria which equilibrate them among cell body, axon, and nerve ending pools.     

Contribution of specific and nonspecific afferent inputs to shaping evoked potentials produced in the cat brain by photic stimulation

Acute and chronic experiments on cats served to show that generation of evoked potentials continued in both the cortex and a number of subcortical structures and that these contained a component matching initial response (except in its reduced amplitude) even after cutting off specific pathways for visual impulse access. This involved severing the optic tract prior to its entry into the lateral geniculate body. Amplitude of primary response decreased less sharply after coagulating the lateral geniculate body, thus preserving a proportion of nonspecific impulse transmission (with pathways via the retino-collicular fibers persisting). Once a major proportion of the nonspecific visual pathways had been destroyed by severing the brachium of the superior colliculi, photic stimulation led to the formation of two-stage evoked potentials with a profile hardly differing from normal. It is presumed that genesis of evoked potentials depends on the quantity rather than the quality of incoming afferents.I. V. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 20, No. 1, pp. 3–7, January–February, 1988.     

Axonal Transport of the Ca2+-Dependent Protein Modulator of 3'':5''-Cyclic-AMP Phosphodiesterase in the Rabbit Visual System

Water-soluble proteins were extracted from individual retinas, optic nerves, combined optic tracts and lateral geniculate bodies, and superior colliculi of rabbits at 1, 3, and 18 days after injection of [3H]leucine into the right eye. The Ca2+-dependent protein modulator of 3':5'-cyclic-AMP phosphodiesterase (calmodulin) was isolated from these samples by a two-step polyacrylamide gel electrophoresis procedure. An analysis of the radioactivity incorporated into the total soluble proteins and the calmodulin revealed that most of the calmodulin was axonally transported at a slow rate (2--4 mm/day) and represented about 0.45% of the total transported soluble protein.     

Effect of destruction of the auditory centers of the brain on the range of echolocation by bats Rhinolophus ferrum-equinum (the large horseshoe-nose bat)

Effect of the auditory cortex, internal geniculate bodies and inferior colliculus removal on echo-ranging detection of immovable wire obstacle by bats has been studied in the behavioural procedure. It is shown that destruction of the auditory cortex results in an irreversible decrease (to 74.6% of control values) in limiting detection range of obstacle by animals. Extensive destructions of internal geniculate bodies decrease limiting detection range only in the first postoperative days. In the case of the inferior colliculi + removal the limiting detection range irreversibly decreases to values equaling 38.5% of control ones. A suggestion that the obtained effects are a consequence of the auditory system sensitivity deterioration in bats is substantiated.     

Functional interrelations of brain structures in the cat during the generation of rhythmic activity. III. Cluster analysis of rhythmic indices

The functional interrelationships of the brain structures of freely moving cats in generation of rhythmic EEG activity during the states of drowsiness and light sleep were evaluated using the claster analysis of mean values indexes of rhythms in different structures as well as correlation coefficients between them in time. It was shown that according these parameters visual cortical areas and lateral geniculate body appeared in different clusters. Lateral geniculate body suggested not to be the only pacemaker of EEG rhythms in visual cortex. The wide convergence of subcortical inputs to the visual cortex and possibility of autonomic generation of EEG rhythms at the cortical level are discussed as putative mechanisms of dissociation of EEG activities in visual cortex and thalamus.     

Early visual experience, learning, and neurochemical plasticity in the rat and the chick.

Rats reared in the dark to 50 days show morphological and biochemical changes in the visual pathway. Light exposure results in elevated incorporation into protein in visual cortex, lateral geniculate and retina. Much of the visual cortex elevation is in a rapidly labelling, rapidly transported neuronal particulate protein. There are concomitant changes in lysosomal and transmitter enzyme activity. In chicks exposed to an imprinting stimulus (a flashing light) there are elevations in RNA polymerase and RNA and protein incorporation in the anterior forebrain roof (a.f.r.) compared with controls. There are changes in adenyl cyclase, cAMP and AChE. Behavioural controls show that although there are general biochemical sequelae of light exposure, the elevation in RNA synthesis in the a.f.r. is not a result of motor, stress or sensory activity, but is correlated with a measure of the learning of the stimulus characteristics. A model for neurochemical correlates of developmental plasticity, learning, and state-dependent transients is discussed.     

Changes in the evoked potentials in the visual and association cortex of the alert cat after changing the order of stimulation of the lateral geniculate body and superior colliculi

In chronic experiments on alert cats the preceding stimulation of the lateral geniculate body (LGB) facilitates the response in the visual cortex to testing stimulation of the anterior colliculi (AC) at all studied delays between stimuli in contrast to the associative cortex where this response has been depressed in intervals of 10-40 ms. In reverse order of stimulation, facilitation of response to testing LGB stimulation is observed at all studied delays in the associative cortex, while in the visual cortex this response is slightly depressed. The obtained data point to the importance of the information coming in the accessory zones of LGB and AC projections and to different informational value of AC inputs to the associative and visual cortices and reciprocal relations between inputs to these cortical areas from LGB and AC.     

First-occupancy model for signal transfer in the brain

In rats and rabbits, optic nerve fibers are asymmetrically decussated. Evoked bioelectric signal/noise power ratio as a function of decussation ratio afferent to the lateral geniculate and thence to visual cortex conforms approximately to what one would expect if the visual system functioned as a parallel coherent detector. Brain mechanisms which can quantitatively account for these findings have not been previously proposed. A theoretical model is proposed to explain observed bioelectric input-output relationships in asymmetrically decussated visual systems and to suggest a general mechanism for signal transfer in the mammalian brain. The model is stochastic and is based upon earlier work concerning coherent neuronal activity. A specific quantitative implication of the model is that, on the average, three ganglion cells in the retina will drive each principal cell in the lateral geniculate. This prediction of the model is verified by recent experimental findings.