Development of neuronal responses in cat posteromedial lateral suprasylvian visual cortex

Exchange assays have often been used to quantitate steroid receptors when endogenous ligands are present; however, there are no reports of their successful application to activated glucocorticoid-Type II receptor complexes. In addition to investigating the reasons for this failure, the present study also examined the effects of progesterone on glucocorticoid dissociation from, and reassociation with unactivated and activated Type II receptors. Molybdate-stabilized brain cytosol from adrenal-ovariectomized mice was incubated with [3H]dexamethasone ( +/- [1H]DEX) for 40 h at 0 degree C. Afterwards free steroid was removed on Sephadex G-25 columns in the presence (unactivated receptors) or absence (activated receptors) of molybdate. Activation, as measured by DNA-cellulose binding, was achieved by incubating molybdate-free cytosol at 22 degrees C for 20 min followed by G-25 filtration in the presence of molybdate. The rates of dissociation and reassociation were then measured by incubating cytosol with [1H]triamcinolone acetonide (TA) or [3H]TA ( +/- [1H]TA) at 12 degrees C. An exchange assay was also employed in which cytosol was incubated first with [1H]DEX for 40 h at 0 degree C followed by bound-free steroid separations and 12 degrees C incubations with [3H]TA ( +/- [1H]TA). Both approaches revealed that even though activation reduced the rate of DEX dissociation from Type II receptors by 40%, it eliminated the ability of the newly unoccupied receptors to rebind glucocorticoid. Adding [1H]progesterone to occupied receptor preparations increased dissociation rate constants by nearly 3-fold, for both unactivated and activated Type II receptors. Since [1H]TA failed to prevent this effect, progesterone appears to act at an allosteric site(s) which cannot be occupied by glucocorticoids. Exchange assays revealed that progesterone-facilitated dissociation increased the rate of glucocorticoid rebinding to unactivated, but not activated Type II receptors. These results suggest that spontaneous and progesterone-facilitated termination of glucocorticoid genomic actions could be mediated by steroid dissociation since unoccupied activated Type II receptors do not rebind agonist steroid.     

Ventral lateral geniculate nucleus afferents to the suprachiasmatic nucleus in the cat

The lateral geniculate complex innervates the hypothalamic suprachiasmatic nucleus (SCN). The location of neurons in the cat ventral lateral geniculate nucleus (vLGN) that give rise to the geniculohypothalamic tract has not been described. In this study, retrogradely labeled neurons were noted throughout the rostrocaudal extent of the medial vLGN following tracer injection into the SCN region. In addition, neuropeptide Y immunoreactive processes were also observed in the vLGN in this same medial zone and in the SCN. The data suggest that the medial zone of the cat vLGN may be homologous to the rodent intergeniculate leaflet (IGL).     

Increased glutamate, GABA and glutamine in lateral geniculate nucleus but not in medial geniculate nucleus caused by visual attention to novelty

This study is concerned with cortico-thalamic neural mechanisms underlying attentional phenomena. Previous results from this laboratory demonstrated that the visual sector of the GABAergic thalamic reticular nucleus is selectively c-fos activated in rats that are naturally paying attention to features of a novel-complex environment, and that this activation is dependent on top-down glutamatergic inputs from the primary visual cortex. By contrast, the acoustic sector of the thalamic reticular nucleus is not activated despite noise generated by exploration and c-fos activation of brainstem acoustic centers (e.g. dorsal cochlear nucleus, inferior colliculus). A prediction of these results is that the levels of the neurotransmitters glutamate and GABA, and the glutamate-related amino acid glutamine, will be increased in the lateral geniculate nucleus (LGN), but not in the medial geniculate nucleus (MGN) of rats that explore a novel-complex environment in comparison to levels of these amino acids in control rats. By means of neurochemical analysis of these amino acids (HPLC) the results of this study confirmed this prediction. The results are consistent with the previously proposed 'focal attention' hypothesis postulating that a focus of attention in the primary visual cortex generates top-down center-surround facilitatory-inhibitory effects on geniculocortical transmission via corticoreticulogeniculate pathways. The results also supports the notion that a main function of corticothalamic pathways to relay thalamic nuclei is attention-dependent modulation of thalamocortical transmission.     

Electrophysiological study of synaptic connections between a transplanted lateral geniculate nucleus and the visual cortex of the host rat

The lateral geniculate nucleus (LGN) of a fetal rat was transplanted to the visual cortex (VC) of a neonatal rat. A current source-density analysis of field potentials and an intracellular study of neuronal responses were conducted in slice preparations by electrical stimulation of transplanted LGN and host VC. The results indicated that synaptic connections were established reciprocally between the transplanted LGN and the host VC.     

Afferent projections to the ventral lateral geniculate nucleus in the cat

Horseradish peroxidase (HRP) injections were made into the dorsal lateral geniculate nucleus (LGN_d) and ventral lateral geniculate nucleus (LGN_v) of the cat in order to define afferent projections to LGN_v. These were found from the superior colliculus, contralateral LGN_v, dorsal median raphe nucleus, locus coeruleus, ipsilateral pretectum, and various portions of visual cortex. While many cortical areas project to LGN_v (17, 18, 19, 21 and lateral suprasylvian), the heaviest input arises from areas 17 and 20. The cell bodies of origin are in layer 5 in contrast to layer 6 which projects to LGN_d.     

Visual pretectal neurons projecting to the dorsal lateral geniculate nucleus and pulvinar nucleus in the cat

We observed morphological subtypes of visual pretectal neurons ascending to the dorsal thalamus, following injections of wheat germ agglutinin conjugated to horseradish peroxidase into the dorsal lateral geniculate nucleus (LGNd) or the pulvinar nucleus. These neurons are composed of fusiform cells and small-sized multipolar cells in the olivary pretectal nucleus, superficial horizontal cells, fusiform cells, small-, medium- and large-sized multipolar cells in the optic tract nucleus, and small- and medium-sized multipolar cells in the posterior pretectal nucleus. When somal size of the neurons projecting to the LGNd was compared to the size of neurons projecting to the pulvinar, the neuronal groups were not identical.     

An analysis of the effect of retinal ganglion cell impulses upon the firing probability of neurons in the dorsal lateral geniculate nucleus of the cat

This study examines the probabilistic way in which LGN cells produce impulses. Simultaneous extracellular recordings were made from a single lateral geniculate nucleus (LGN) relay cell and the one retinal cell that triggered most of its impulses during vigorous responses. Impulses in the retinal train were classified as 'successful' in triggering an LGN impulse, or 'unsuccessful'. Similarly, the impulses in the LGN train were either 'Triggered' by a successful retinal impulse, or were 'Anonymous'. These impulses delimited various intervals whose distributions were compared to the relevant distribution of all intervals to determine whether short or long intervals tended to dominate in each case. Intervals between unsuccessful and successful impulses tended to be shorter than other retinal intervals, with their probability declining exponentially with duration. These data imply a decaying excitation produced by each impulse, but with a short refractory period following each Triggered impulse. Short intervals between Anonymous impulses were relatively common; Anonymous impulses thus lack the same refractoriness and tend to occur in bursts. The exponential excitation following an unsuccessful retinal impulse also facilitates Anonymous impulses, while Anonymous impulses (during visual stimulation) render the LGN slightly refractory for subsequent retinal impulses.     

Development of the projections from the dorsal lateral geniculate nucleus to the lateral suprasylvian visual area of cortex in the cat.

In the study reported in the preceding paper, we used retrograde labeling methods to show that the enhanced projection from the thalamus to the posteromedial lateral suprasylvian (PMLS) visual area of cortex that is present in adult cats following neonatal visual cortex damage arises at least partly from surviving neurons in the dorsal lateral geniculate nucleus (LGN). In the C layers of the LGN, many more cells than normal are retrogradely labeled by horseradish peroxidase (HRP) injected into PMLS cortex ipsilateral to a visual cortex lesion. In addition, retrogradely labeled cells are found in the A layers, which normally have no projection to PMLS cortex in adult cats. The purpose of the present study was to investigate the mechanisms of this enhanced projection by examining the normal development of projections from the thalamus, especially the LGN, to PMLS cortex. Injections of HRP were made into PMLS cortex on the day of birth or at 1, 2, 4, or 8 weeks of age. Retrogradely labeled neurons were present in the lateral posterior nucleus, posterior nucleus of Rioch, pulvinar, and medial interlaminar nucleus, as well as in the LGN, at all ages studied. Within the LGN of the youngest kittens, a small number of retrogradely labeled cells was present in the interlaminar zones and among the cells in the A layers that border these zones. Such labeled cells were virtually absent by 8 weeks of age, and they are not found in normal adult cats. Sparse retrograde labeling of C-layer neurons also was present in newborn kittens.(ABSTRACT TRUNCATED AT 250 WORDS)     

Differential development of cation-chloride cotransporters and Cl- homeostasis contributes to differential GABAergic actions between developing rat visual cortex and dorsal lateral geniculate nucleus

A recent study suggested that gamma-aminobutyric acid (GABA) plays differential roles in activity-dependent plasticity between the visual cortex (VC) and the dorsal lateral geniculate nucleus (dLGN). In the present study, to investigate differential GABAergic functions in postnatal visual system development, the development of [Cl(-)](i), cation-Cl(-) cotransporter expression, and the [Ca(2+)](i) responses evoked by GABA were compared between VC and dLGN during the early stages of development. Using rat brain slices from postnatal days (P) 0-17, GABA-evoked [Ca(2+)](i) responses and resting [Cl(-)](i) were measured by means of optical imaging of Ca(2+) and Cl(-), respectively. Changes in the expression of cation-Cl(-) cotransporters (viz. the outwardly-directed K(+)-Cl(-) cotransporter, KCC2, and the inwardly-directed Na(+),K(+)-2Cl(-) cotransporter, NKCC1) were examined in VC and dLGN by in situ hybridization. At birth, the excitatory actions of GABA were powerful in VC, but missing in dLGN (as indicated by neuronal [Ca(2+)](i) transients), and the resting [Cl(-)](i) was significantly higher in VC than in dLGN. Signals for KCC2 mRNA expression were significantly higher in dLGN than in VC at P0. This suggests that extrusion of Cl(-) from neurons is stronger in dLGN than in VC at P0, so that a GABAergic excitatory effect was not observed in dLGN because of more negative equilibrium potential for Cl(-). The present study indicates clear differences in the molecular and physiological bases of Cl(-) homeostasis and GABA actions between the developing VC and dLGN. Such differential GABAergic actions may underlie the distinct mechanisms involved in VC and dLGN development within the visual system.     

Visual function in the ventral lateral geniculate nucleus of the cat

The visual receptive fields of 293 single units in the ventral lateral geniculate nucleus of the cat were studied. In addition to the wide variety of types described by others, a group of units responding differentially to color was identified that included units responding particularly to blue and others with opponent color properties. Some units with spontaneous firing and without definite visual receptive fields were inhibited by stimulation of the optic chiasm (OX). A study of latency of firing to OX stimulation suggested that these cells were driven by retinal ganglion cells of the W type. One-third of all units studied were binocularly driven.     

Ultrastructure of synapses from the A-laminae of the lateral geniculate nucleus in layer IV of the cat striate cortex

The morphology of synapses in layer IV of the cat striate cortex was studied by electron microscope (EM) autoradiography of serial sections following injection of tritiated amino acids into the lateral geniculate nucleus. Of the terminals in the neuropil, 22% had 2 or more silver grains in 10 successive sections and were labeled at 8-80 times the background level. These terminals were considered to be specifically labeled and to be derived from the lateral geniculate. Two forms of geniculate synapse were observed. One had medium-size, round vesicles and a modest postsynaptic asymmetry (RA); the other had smaller, pleomorphic vesicles and hardly any postsynaptic opacity; that is, it appeared symmetrical (PS). The geniculate RA terminals were presynaptic to dendritic spines, fine processes, and cell bodies; the geniculate PS terminals were presynaptic to dendrites and cell bodies but not to spines. The possible sources of geniculate PS terminals are discussed.     

Synaptic delay in the lateral geniculate nucleus of the cat

Unitary, presynaptic spike potentials were observed in single cell recordings from the dorsal lateral geniculate nucleus of the cat. In 11 cells, spontaneous S potentials (extracellularly recorded excitatory postsynaptic potentials) were preceded at a fixed interval by a small wave (the 'T' potential). In another 14 cells, a T potential, although not detected in single traces, was revealed by averaging 20-100 samples synchronized to the peak of the S potential. Provided the field response was not too large a T potential could also be detected in the response to a stimulus to the optic nerve. The T potential would appear to be the spike potential of the afferent optic axon which is excitatory to the geniculate cell because it precedes the S potential at a very exact interval and also follows the corresponding retinal ganglion cell spike at a very exact interval and because the interval between T potential and S potential is reversibly decreased by cooling with a temperature coefficient characteristic of synapses. T potentials ranged in amplitude from 8 to 134 microV and were all predominantly positive-going suggesting a failure of the nerve impulse to invade fully the terminals of the optic nerve. The time from the positive peak of the T potential to the start of the S potentials was taken as a good measure of the synaptic delay. The T-S interval averaged 0.29 ms (+/- 0.045 ms S.D.).     

Lack of visual suppression in the rabbit lateral geniculate nucleus during blink reflex

Stimulation of the supraorbital branch of the trigeminal nerve (SO) elicited eye blinks in the rabbit, but did not decrease the amplitude of visual cortical evoked potential from stimulation of the optic chiasm (OX). In addition, the SO stimulation neither induced an inhibitory postsynaptic potential (IPSP) in LGN cells, nor activated inhibitory interneurons in the thalamic reticular nucleus (TRN), which proved to mediate both recurrent inhibition and saccadic suppression in the dorsal lateral geniculate nucleus (LGN). All these indicate that there is no visual suppression in the rabbit LGN during blink reflex.     

Structural development of the lateral geniculate nucleus and visual cortex in monkey and man

This study concerns the development of the primary visual pathway of the primate. The lateral geniculate nucleus (LGN) is the principal thalamic relay to the visual cortex (area 17), and its neurons have similar morphological characteristics in both monkey and man, as identified by Golgi impregnation. The commonest neuron is the multipolar with a radiate or tufted dendritic tree; next is the bipolar neuron with two or three diametrically opposed dendritic trunks. Less frequent are neurons with beaded dendrites and others with fine, axon-like dendritic processes, possibly interneurons. The dendritic tree of all neurons remains generally within a lamina, but some dendrites cross interlaminar zones. LGN neurons are identifiable before birth and differ from their adult form by the presence of immature features, especially numerous dendritic and somatic spines, most frequent at birth in monkeys and at about 4 months postnatally in man. They disappear almost completely by 3 months in monkeys and 9 months in man. The human LGN has reached its ‘adult’ volume by this age.Two stages in the development of the human area 17 can be defined. The first is marked by a rapid growth to its ‘adult’ volume by about 4 months, and by intense synaptogenesis beginning in the foetus and reaching a maximum around 8 months. The second stage is one of stabilization in the volume of area 17 and loss of synapses to reach ‘adult’ synaptic density around 11 years, at about 60% of the maximum values.The formation of transitory morphological features in the first weeks or months of life coincides with a period of visual plasticity in infant monkeys and humans. Our observations can be correlated with experimental evidence for visual development in monkeys and with clinical evaluation of visual activity during the human preverbal stage, a period of great importance in the establishment of visual acuity, of stereopsis and of oculomotor function, all very sensitive to the numerous forms of visual deprivation.     

Projection of brain stem neurons to the perigeniculate nucleus and the lateral geniculate nucleus in the cat

Small horseradish peroxidase injections in the perigeniculate nucleus (PGN) or the lateral geniculate nucleus (LGN) gave retrograde labeling of many cells in the pontomesencephalic reticular formation (RF), the nuclei raphe dorsalis and centralis linearis, locus coeruleus, nucleus of the optic tract and nucleus parabigeminalis. Antidromic stimulation was used to identify neurons in the RF projecting to the PGN-LGN complex. Threshold mapping through the PGN and the LGN shows separate projection from the reticular formation to the PGN and the LGN.     

The morphology of corticofugal axons to the dorsal lateral geniculate nucleus in the cat

The structural features of corticogeniculate axons were studied in adult cats after labeling them with horseradish peroxidase (HRP). Injections of HRP into the optic radiations near the dorsal lateral geniculate nucleus result in Golgi-like filling of both geniculate relay neurons and corticogeniculate axons. In the present material at least two main types of axons could be defined. The most common type is called the type I axon because it so closely resembles the type I axons described by Guillery ('66, '67) in Golgi preparations. These fine axons have smooth surfaces and consistent fiber diameter. Most terminal swellings are at the ends of short collateral branches and these swellings form asymmetric synaptic contacts onto small and medium-sized dendrites. Type I axons typically innervate more than one lamina as well as interlaminar zones and they clearly arise from the cerebral cortex. The second type of axon is called the beaded axon because of its numerous swellings, en passant. These swellings frequently are larger than those on type I axons and they differ from previously described corticogeniculate axon terminals in their ultrastructural features. That is, their synaptic contacts appear symmetrical and they form axosomatic contacts. Because of these differences, the possibility that beaded axons are of subcortical origin, particularly from the perigeniculate nucleus, is discussed. When type I axons and geniculate relay neurons are filled in the same region of the nucleus it is possible to identify probable sites of synaptic contact by using the light microscope. Such analyses indicate that corticogeniculate axons synapse directly onto relay cells, primarily on peripheral dendritic branches. Further, it appears that single axons contact many geniculate neurons and that single neurons are contacted by many axons.     

Local excitability in the superior colliculus influences evoked responses of lateral geniculate cells in rabbits

In anesthetized, immobilized rabbits recordings were made simultaneously from cells in the Lateral Geniculate Nucleus (CGL) and Superior Colliculus (CS), in order to study how the CS influences the CGL. The experimental protocol consisted of three steps. In the initial step (first control) the light stimulus was triggered electronically. In the second step (Test), the same stimulus was triggered by a spontaneous spike arising from a collicular cell. Thus the stimulus presentation was time-locked to collicular endogenous activity. The third step was the same as the first and constituted a second control. The frequencies of stimulus application were gated to be approximately the same. The results indicated that the CS exerts two separate effects on CGL units. In 37 pairs (26%), conditioning the stimulus presentation to collicular firing produced a significant enhancement of geniculate responses. In 24 pairs (17%), the geniculate responses declined. In 82 pairs (57%), no significant influence was noted. The colliculo-geniculate inlfuence is transient. The effects peaked between 100 to 200 msec after the collicular spike and returned to their control levels within 300 msec. Collicular cells producing a decline were encountered mostly in the ventral part of the stratum griseum superficiale, and the stratum opticum, whereas collicular cells that were related to an increased geniculate response were more frequently found dorsally. Increments were more pronounced if the distance (D) between receptive fields was short (0° < D < 40°) or if the collicular and geniculate fields were far apart (120° < D < 180°). The decrement effect was attenuated as the distance separating the two receptive fields. This study suggests that the superior colliculus is capable of generating an internal signal powerful enough to modulate at the geniculate nucleus the visual message conveyed toward the visual cortex. A possible role of the CS in the initiation of the corollary discharge is briefly discussed.     

Origin of butyrylcholinesterase in the lateral geniculate nucleus of tree shrew

We examined the distribution and possible origins of pseudocholinesterase activity within the lateral geniculate nucleus (LGN) of the tree shrew. Butyrylcholinesterase (BuChE) activity was spread diffusely throughout the LGN and not localized to neuronal perikaryon. Lesions of the LGN eliminated this BuChE activity while not affecting acetylcholinesterase (AChE) activity; however, removal of retinal input by unilateral ocular enucleations failed to affect the BuChE activity within the denervated layers of the LGN. This lack of effect suggests that, unlike the macaque monkey, retinal terminals within the LGN of tree shrew are not the source of BuChE. Thus, in the tree shrew LGN it appears that BuChE is not metabolically related to or dependent upon AChE nor does it originate from retinal sources, but rather BuChE appears to represent an enzyme that is endogenous to the LGN.