Axon collaterals of mossy fibers from the pontine nucleus in the cerebellar dentate nucleus.

1. Single axons of pontine nucleus neurons (PN axons) receiving cerebral input were stained intra-axonally with horseradish peroxidase (HRP) in the cerebellum of cats. The axonal trajectory of single PN axons was reconstructed from serial sections of the cerebellum and the brain stem. 2. Axons were penetrated in the white matter near the dentate nucleus, and, after electrophysiological identification, PN axons were injected iontophoretically with HRP. The identification criteria for the PN axons were 1) their direct responses to stimulation of the contralateral pontine nucleus (PN), 2) their synaptic activation from the contralateral cerebral cortex, and 3) the decrease in threshold for evoking direct spikes in stimulation of the PN by conditioning stimuli applied in the cerebral cortex. 3. Two hundred thirty-three axons were electrophysiologically identified as PN axons receiving the input from the cerebral cortex. Ninety-six of them were stained successfully with HRP, and reconstructions were made from 40 well-stained PN axons. All of them gave rise to mossy fibers and terminated in the granular layer of the cerebellar cortex as typical mossy fiber rosettes. Out of these, 22 gave axon collaterals to the dentate nucleus. Virtually all of the axon branches observed in the dentate nucleus were axon collaterals of mossy fibers from the PN to the cerebellar cortex. In 7 of these 22 PN axons, cell bodies were retrogradely labeled with HRP, and all of them were found in the contralateral PN. 4. The stained-stem axons arising from the PN ran medially in the pons, crossed the midline, and then ascended dorsocaudally in the branchium pontis. After passing in the white matter anterior to or lateral to the dentate nucleus, they entered into the cerebellar cortex. On their way, one to three axon collaterals were given off from parent axons to the dentate nucleus. The diameter of these collaterals was very thin (mean, 0.6 microns), compared with the large diameter of the parent axons (mean, 2.1 microns). 5. Some axon collaterals were very simple and had only one terminal branch with or without short branchlets, whereas others were more complex, and single axon collaterals ramified before forming a terminal arborization. Axon collaterals of single PN axons mainly spread mediolaterally or dorsoventrally in the frontal plane but had a very narrow rostrocaudal extension. 6. Terminal branches usually bore swellings en passant along their length and one terminal swelling at their end. The number of swellings per axon collateral ranged 23-180 (116 +/- 52, mean +/- SD).(ABSTRACT TRUNCATED AT 400 WORDS)     

Morphology of physiologically identified retinogeniculate X- and Y-axons in the cat

1. We studied the morphology of individual, physiologically identified retinogeniculate axons in normal adult cats. The axons were recorded in the lateral geniculate nucleus or in the subjacent optic tract, characterized as X or Y by physiological criteria, penetrated, and injected with horseradish peroxidase. With subsequent application of appropriate histochemistry, the enzyme provides a complete label of the terminal arbors and parent trunks for morphological analysis. We have recovered for such analysis 26 X- and 25 Y-axons; of these, 14 X- and 12 Y-axons were studied in detail. 2. Within the optic tract, the parent trunk of every X-axon is located closer to the lateral geniculate nucleus and thus further from the pial surface than that of every Y-axon. This probably reflects the earlier development of X- than of Y-axons. Furthermore, the parent axon trunks of the X-axons are noticeably thinner than are those of the Y-axons. Every retinogeniculate X- and Y-axon in our sample branches within the optic tract. One of these branches heads dorsally to innervate the lateral geniculate nucleus and one heads medially and rostrally toward the midbrain, although none of these labeled axons were traced to a terminal arbor beyond the lateral geniculate nucleus. For Y-axons, all branches are of comparable diameter, but for X-axons, the branch heading toward the lateral geniculate nucleus is always noticeably thicker than is the branch directed toward the midbrain. 3. Every retinogeniculate X- and Y-axon produces the greatest portion of its terminal arbor in lamina A (if from the contralateral retina) or A1 (if from the ipsilateral retina). These arbors typically extend across most of the lamina along a projection line. Not a single terminal bouton from any axon was found in the inappropriate lamina A or A1 (i.e., in lamina A for ipsilaterally projecting axons or in lamina A1 for contralaterally projecting ones). Occasionally, an X-axon also innervates the medial interlaminar nucleus, and even more rarely does an X-axon innervate the C-laminae. In contrast, nearly all Y-axons from the contralateral retina branch to innervate part of the C-laminae (probably lamina C), and most from either retina also innervate the medial interlaminar nucleus. Although these details imply considerable variation in the overall pattern of retinogeniculate innervation for both X- and Y-axons, we found no physiological properties to correlate with this variation.(ABSTRACT TRUNCATED AT 400 WORDS)     

Intracellular recordings from binocularly activated cells in the cat's dorsal lateral geniculate nucleus

Five binocularly activated cells near the interlaminar layers of the dorsal lateral geniculate nucleus have been studied with intracellular recording techniques. Four neurons were relay cells and antidromically activated from the visual cortex. They received monosynaptic excitation and disynaptic inhibition from Y type retinal ganglion cells in both eyes and disynaptic recurrent inhibition. The fifth cell was similar to perigeniculate neurons. It received disynaptic excitation from retinal ganglion cells in both eyes and monosynaptic excitation from antidromically activated relay cell axons. It was also inhibited from all these sources after an additional synaptic delay. The cell had a large receptive field, about twice the center size of neighboring relay cells, and gave on-off responses from the entire response area. Such displaced perigeniculate like cells may explain why relay cells issue occasional axon collaterals within the dorsal lateral geniculate nucleus.     

Alpha and beta cells projecting from retina to lamina A of the lateral geniculate nucleus in normal cats,monocularly deprived cats,and young kittens

Summary We strictly limited small injection of horseradish peroxidase (HRP) to lamina A of the lateral geniculate nucleus of cats. This was done to label retrogradely only the alpha (Y) and beta (X) classes of retinal ganglion cell. Eighty-six such injections at a range of matched eccentricities were made bilaterally in 9 normal adult cats, 7 cats reared from birth to adulthood with monocular lid suture, and 9 normal kittens at 4 weeks of age; 5348 alpha and beta cells were retrogradely labeled from these injections. Quantitative measurements were made from these labeled cells and compared among 4 experimental conditions, these being normal adult retinas, the nondeprived and deprived retinas of lid sutured cats, and the retinas of kittens. Each injection led to a similar relative ratio of labeled alpha and beta cells (typically 5–15% alpha cells) that did not differ significantly among the experimental conditions, but further analysis suggested a slight dimunition of labeled alpha cells in deprived retinas. Because the larger arbors of retinogeniculate Y axons are more likely to penetrate small geniculate HRP injection sites from eccentric locations than would be the case for the more restricted arbors of X axons, a normal tendency resulted for the peripheral halo of zones of retrograde labeling to be dominated by alpha cells. Thus a more accurate reflection of the relative numbers of labeled alpha and beta cells would result from considering only the core of zones of retrograde labeling. When this is done, deprived retinas exhibited relatively fewer labeled alpha cells than did normal, nondeprived, or kitten retinas. This may relate to prior observations (Sur et al. 1982) that abnormally few Y axons from the derpived retina innervate lamina A. No statistically significant differences in alpha or beta cell size were seen among normal, nondeprived, and deprived retinas, although both of these cell types in the kittens were equally smaller than their normal adult counterparts. This is particularly interesting in view of the postnatal growth of retinogeniculate axon arbors (Sur et al. 1984). The results are not surprising for alpha cells, since retinogeniculate Y axon arbors grow considerably after 4 weeks of age, but they are surprising for beta cells, since retinogeniculate arbors of X axons decrease after 4 weeks of age. This suggests no clear, general relationship between soma size and the extent of a cell's axonal arbor. Overall, these results suggest that no dramatic abnormalities due to rearing with monocular suture are evident at the level of the retina, although subtle effects can be demonstrated there (see also Leventhal and Hirsch 1983). The most peripheral site in the visual system at which such dramatic effects have been documented thus seems to be at the level of retinogeniculate innervation.     

Identification of X versus Y properties for interneurons in the A-laminae of the cat's lateral geniculate nucleus

Summary Roughly 25% of the neurons in the A-laminae of the cat's lateral geniculate nucleus are local interneurons, while the remaining 75% are relay cells that project to the visual cortex. The interneurons form the focus of our study. The relay cells are either X or Y cells and are thereby integral links in the parallel and independent retino-geniculo-cortical X and Y pathways. Little is known about the response properties of interneurons, largely because it is difficult to identify them clearly during electrophysiological recording. However, they can be identified by morphological criteria. We thus studied their response properties by recording intracellularly from geniculate neurons to characterize them and then injecting them with horseradish peroxidase (HRP); the HRP labeling subsequently allowed us to distinguish relay cells from interneurons. In this manner, we studied 171 relay cells (83 X and 88 Y) and 15 interneurons. The response properties tested for each of the interneurons were indistinguishable from those of the relay X cells. We conclude that these interneurons are directly innervated by retinogeniculate X axons and are firmly embedded in the X pathway. We found no evidence for inter-neurons in the Y pathway.     

Corticofugal axons in the lateral geniculate nucleus of the cat

Summary Injections of horseradish peroxidase (HRP) were made into the optic radiations just above the lateral geniculate nucleus of four cats to trace the anterograde filling of corticofugal axons terminating in the perigeniculate and lateral geniculate nuclei. The different types of axons were classified according to axonal diameter and the morphology of the terminal appendages. Judging from their morphological organization we suggest that the corticofugal axons are, in the main, slowly conducting and that they have a restricted terminal distribution which extends, however, to a multiplicity of levels in both perigeniculate and lateral geniculate nuclei. These morphological characteristics may have physiological implications in determining the role of the corticofugal pathway.     

Feedback inhibition in the cat's lateral geniculate nucleus

Feedback inhibition is generally believed to be a ubiquitous feature of brain circuitry, but few specific instances have been documented. An example in cats is the supposed feedback circuit involving relay cells of the lateral geniculate nucleus and cells of the perigeniculate nucleus (a part of the thalamic reticular nucleus): geniculate relay cells innervate the perigeniculate nucleus, which, in turn, provides an inhibitory, GABAergic projection back to the lateral geniculate nucleus. However, feedback inhibition at the single-cell level requires that a given perigeniculate cell project back onto the same geniculate relay cell that innervates it. We probed for this in an in vitro slice preparation of the cat's lateral geniculate nucleus. We evoked a single action potential in a geniculate cell via a brief, depolarizing pulse delivered through an intracellular recording electrode and looked for any evoked hyperpolarizations. For 6 of the 36 geniculate cells tested, we observed a long-lasting hyperpolarization after the action potential, and much of this was eliminated by application of bicuculline, suggesting synaptically activated inhibitory postsynaptic potentials. We interpreted this to be clear evidence that a given neuron may inhibit itself via circuitry mediating feedback inhibition in the cat's lateral geniculate nucleus.Shanghai Brain Research Institute, Shanghai, People's Republic of China 200031     

The morphology of neurons in trigeminal nucleus oralis projecting to the medullary dorsal horn (trigeminal nucleus caudalis): a retrograde horseradish peroxidase and Golgi study

This study demonstrates that trigeminal nucleus oralis, the most rostral subdivision of the spinal trigeminal nucleus, contains four morphologically distinct types of small neurons which project to the medullary dorsal horn (trigeminal nucleus caudalis) via descending intratrigeminal pathways. Using the retrograde transport of horseradish peroxidase following injections in the medullary dorsal horn, labeled small neurons with cell bodies ranging from 8-15 microns in diameter are found principally in the ventrolateral portion of the trigeminal nucleus oralis. Most neurons are labeled ipsilaterally throughout the entire rostrocaudal extent of the ventrolateral portion of the trigeminal nucleus oralis, but a few cells are also labeled contralaterally. From this aspect of the present study it can be concluded that a specific portion of the trigeminal nucleus oralis, i.e. the ventrolateral part, contains numerous small neurons which send descending projections to the medullary dorsal horn that could affect synaptic activity there. Utilizing both the methods of Golgi and retrograde horseradish peroxidase labeling four distinct types of small descending medullary dorsal horn projection neurons can be distinguished in the ventrolateral portion of the trigeminal nucleus oralis on the basis of their morphology and the distribution of their axons and dendrites. All four neuronal cell types are present throughout the entire rostrocaudal extent of the trigeminal nucleus oralis. Type I neurons are the most frequently labeled descending medullary dorsal horn projection neurons. They are concentrated in the medial 500-550 microns of the ventrolateral portion of the trigeminal nucleus oralis and display dendritic trees which occupy spherical domains approaching 300 microns in diameter. The unmyelinated axons of many of these cells arise either directly from the cell body or a primary dendrite and give rise to a single collateral within 50 microns of their site of origin. This collateral generates a fine axonal plexus within a portion of the dendritic arbor of the parent cell while the parent axon, without branching further, travels a short distance in the ventrolateral portion of the trigeminal nucleus oralis and enters a deep axon bundle. Type II neurons are the second most frequently labeled descending medullary dorsal horn projection neuron. They generate medial and lateral dendritic arbors which together span nearly the entire medial 500-550 microns of the ventrolateral portion of the trigeminal nucleus oralis. An unmyelinated axon emerges from the cell body and within 10-30 microns of its origin gives rise to two collaterals.(ABSTRACT TRUNCATED AT 400 WORDS)     

Ultrastructural identification of synaptic terminals from cortical axons and from collateral axons of geniculo-cortical relay cells in the perigeniculate nucleus of the cat

Summary Electron microscopic analysis of sections of the perigeniculate nucleus (PGN) of the cat processed with horseradish peroxidase (HRP) histochemistry after massive injections of this enzyme in the visual cortex showed two types of synaptic terminals labeled with HRP reaction products. One type (RLD terminals) is characterized by round synaptic vesicles, large size, dark mitochondria and asymmetrical synaptic contacts with somata and dendrites. The second type (RSD terminals) is characterized by round synaptic vesicles, small size, dark mitochondria and asymmetrical synaptic contacts with dendrites. The HRP + RSD terminals, which were also found in the dorsal lateral geniculate nucleus (LGN), are interpreted as terminals of cortical origin both in the PGN and LGN, since previous studies have identified cortical terminals as being of RSD type in the LGN and in other thalamic nuclei. The HRP + RLD terminals are interpreted as synaptic terminals of collaterals axons of geniculo-cortical relay cells in the PGN labeled by retrograde transport of HRP from the cortex. In addition, in semithin and ultrathin sections somata in the PGN were never found labeled with HRP products indicating the absence of a PGN projection to the visual cortex.     

A GABAergic projection from the pretectum to the dorsal lateral geniculate nucleus in the cat.

To study the projection from the pretectum to the lateral geniculate nucleus, we placed wheat-germ agglutinin conjugated to horseradish peroxidase into the lateral geniculate nuclei of six cats, allowed this marker to be retrogradely transported by afferent axons to their parent somata in the pretectum, and revealed the label in these cells with stabilized tetramethylbenzidine histochemistry. In three cases we made large pressure injections that completely infiltrated the lateral geniculate nucleus and extended into neighboring thalamic nuclei; in the other three we made smaller iontophoretic injections largely confined to the A- and C-laminae of the lateral geniculate nucleus. In both types of injection we found labeled pretectal cells mainly in the nucleus of the optic tract but also found some cells labeled in the olivary pretectal nucleus and the posterior pretectal nucleus. After one of the larger injections we analysed both sides of the pretectum and found that 11% of the labeled cells were located contralaterally and were distributed in the same three nuclei. We analysed only the ipsilateral side in the remaining five cats. In those five experiments we also immunohistochemically stained the pretectal sections with an antibody directed against the neurotransmitter, GABA. Of the retrogradely labeled pretectal cells, 40% were also labeled for GABA, and those were similar in soma size (350 microns 2 in cross-sectional area) to those labeled only with the retrograde marker (331 microns 2). GABA-positive cells not labeled by retrograde transport were smaller (246 microns 2) than either of these other cells populations. These results indicate that at least 40% of the cells involved in the projection from the pretectum to the lateral geniculate nucleus are GABAergic. We suggest that this extrathalamic projection may serve to inhibit thalamic GABAergic cells. This, in turn, would disinhibit geniculate relay cells, thereby facilitating the geniculate relay of retinal information to cortex.     

Development of terminal arbors of retino-geniculate axons in the kitten—I. Light microscopical observations

The maturation of terminal arbors of retino-geniculate axons was studied in normal kittens from 1 to 8 postnatal weeks. Horseradish peroxidase injected into the optic tract rostral to the lateral geniculate nucleus gave a dense fill of cut axons and their terminals, resembling results obtained by the Golgi methods. At 1 and 2 weeks postnatal, the overall size and extent of axon arbors is not significantly different than in the adult. However, terminal branches of axon arbors at this age give rise to high variable endings. They terminate in finely divided sprays of finger-like extensions and filopodia bearing small spikes rather than the characteristic clusters and strands of crenulated terminals of adult axons. The bases of these sprays are broad and irregular in contour, with foliate growth-cone-like structures occurring at the ends of some branches. At 3 weeks postnatal, terminal swellings become thicker and more crenulated in contour. By 5 to 6 weeks, axon arbors have adult-like terminals with respect to their size, shape and arrangement, although entire branches may still be immature with irregular terminal swellings. Small slightly indented terminals are also seen for the first time. By 8 weeks, axon arbors are generally mature, but occasionally have a bizarre and immature arrangement of fine extensions or growth-cone-like tips.Although these observations do not establish whether reduction of branches or of terminals takes place during postnatal maturation, they demonstrate that kitten retino-geniculate axon terminal arbors are highly immature and undergo considerable changes during the period optimal for induction of sprouting by eye enucleation. The morphogenetic maturation of terminal branches that begins at 3 weeks also marks a decline in their sprouting capacity, even though remodeling of terminals is not complete until after 8 weeks of age.     

Induction of somatic sensory inputs to the lateral geniculate nucleus in congenitally blind mice and in phenotypically normal mice.

The incidence of aberrant innervation of the lateral geniculate nucleus by ascending somatic sensory axons was examined following injections of wheat germ agglutinin conjugated with horseradish peroxidase into the dorsal column nuclei of adult mice which were: (1) normal; (2) normal, but bilaterally enucleated on the day of birth; (3) normal, but received a large unilateral lesion of the rostral cortex on the day of birth; (4) normal, bilaterally enucleated, as well as unilaterally lesioned in the rostral cortex on the day of birth; (5) homozygous for an ocular retardation mutation (orj/orj); or (6) homozygous for the orj mutation and received a large unilateral lesion of the rostral cerebral cortex on the day of birth. In the phenotypically normal animals which were untreated, no somatic sensory inputs enter into the dorsal lateral geniculate nucleus. A few labeled axons enter into and arborize within the dorsal lateral geniculate nucleus in normal animals which received bilateral enucleations or unilateral rostral cortical lesions on the day of birth. However, in congenitally blind animals and in phenotypically normal animals which received bilateral enucleations as well as unilateral rostral cortical lesions on the day of birth, a significant number of labeled axons enter into and arborize within the dorsal lateral geniculate nucleus. Among all these experimental groups, the densest innervation of the lateral geniculate nucleus occurred in congenitally blind animals which received rostral cortical lesions on the day of birth. In these, robust arborizations of labeled somatic sensory axons occupy a substantial extent of the lateral geniculate nucleus. These results not only demonstrate that ascending somatic sensory axons can be rerouted to the lateral geniculate nucleus, but also indicate that the ability of a thalamic afferent pathway to undergo extensive reorganization and to innervate inappropriate thalamic targets following early perturbations is not unique to the retinal projection (in which this has previously been demonstrated), and may be a more general characteristic of the major thalamic afferent systems.     

The GABA-immunoreactive neurons in the interlaminar regions of the cat lateral geniculate nucleus: light and electron microscopic observations

Summary GABA-immunoreactive cells located in the interlaminar zone between the A and A1 laminae of cat LGN were studied at the LM and EM levels. The mean perikaryal size of these neurons was larger than that of GABA-immunoreactive cells in the A-laminae of LGN. Interlaminar GABA+ cells examined in plastic semithin sections of LGN after massive injections of HRP in the striate and extrastriate visual cortex were not retrogradely labeled with reaction products (as previously reported for the GABA+ cells in the laminar regions of LGN) suggesting that these cells do not project to the visual cortex. Serial EM analysis of two partially reconstructed interlaminar GABA+ cells showed that they receive synaptic inputs from RLD terminals of axon collaterals of geniculo-cortical relay cells, from cortical (RSD) terminals, from inhibitory (F) axon terminals, and from other undetermined terminals, but not from retinal (RLP) axon terminals. These data suggest that the GABAergic cells in the interlaminar zones of LGN participate as interneurons in recurrent inhibitory circuits in LGN. The synaptic inputs to these cells and ultrastructural features, notably somatic spines and dendrites oriented predominantly orthogonal to the projection lines in LGN, are similar to those of neurons of the perigeniculate nucleus.     

Identification of geniculo-tectal relay neurons in the rat's ventral lateral geniculate nucleus

Summary In the rat's ventral lateral geniculate nucleus (vLGN), geniculo-tectal relay neurons (GTR-neurons) could be identified by the retrograde transport of horseradish peroxidase (HRP) after injection in the superior colliculus (SC). GTR-neurons correspond to class III cells described by Brauer and Schober (1973) in Golgi preparations of the rat's vLGN. The distribution of GTR-neurons is restricted to the lateral subnucleus of vLGN. According to Swanson et al. (1974), the axons of these cells terminate in lower Stratum griseum superficiale and in Stratum opticum, Stratum griseum intermedium and Stratum album intermedium of SC.The GTR-neurons are characterized by very thick and long proximal dendritic segments which have a smooth surface. Dendrites branch preponderantly in their distal regions and only in this part form many multiform protrusions. There is some evidence that retinal axons terminate on these dendritic surface structures. The supposed differences in the afferent patterns between GTR-neurons in the vLGN and geniculo-cortical relay neurons in the dorsal lateral geniculate nucleus are discussed.Sponsored by a grant of the Ministry of Science and Technology of the GDR     

Effects of monocular deprivation on geniculocortical pathways in the owl monkey, Aotus trivirgatus

As has been shown for a number of other mammals, monocular eyelid suture in the newborn owl monkey, a New World primate, is followed by a failure of neurons in the deprived layers of the lateral geniculate nucleus to reach normal size in the adult animal. This failure appears to be almost completely confined to relay cells, identified by horseradish peroxidase (HRP) injections into cortex, rather than interneurons. Similar proportions of neurons in both the deprived and non-deprived layers are labeled after HRP injections, suggesting that relay cells do not lose all cortical connections as a result of visual deprivation.     

The thalamic reticular nucleus of the adult rat: experimental anatomical studies

Summary The thalamic reticular nucleus (TRN) is a sheet-like nucleus partially enclosing the dorsolateral and anterior aspects of the thalamus and traversed by the thalamo-cortical and cortico-thalamic fibre systems. This paper describes the cellular and synaptic organization of the TRN in adult albino rats on the basis of LM and EM studies of normal animals and experimental animals with injections of horseradish peroxidase (HRP) and/or lesions in various parts of the brain. Particular attention was paid to the dorso-caudal part of the TRN, which establishes connections with visual centres.LM-HRP preparations show that the neurons of TRN project only to ipsilateral dorsal thalamus; no labelled cell bodies were found in TRN after injections into the cortex or any part of the brain stem caudal to the thalamus. Small injections into dorsal thalamus result in a small cluster of labelled neurons and an associated patch of terminal label in TRN. The dorso-caudal part of the nucleus projects to the dorsal lateral geniculate nucleus, the ventro-caudal part to the medial geniculate nucleus and a large part of the nucleus anterior to the areas associated with the geniculate nuclei projects to the ventrobasal nucleus. No evidence was found for a widespread distribution of reticulo-thalamic axons and the connections between TRN and the dorsal lateral geniculate nucleus and between TRN and the ventrobasal nucleus show a fine-grain topographical organization with more rostral and dorsal parts of TRN projecting to more rostral and dorsal parts of the dorsal lateral geniculate and ventrobasal nuclei.The neurons of TRN are variable in size (range of somal diametersc. 10–20 m), shape (cell bodies are most commonly ellipsoidal) and dendritic morphology (bitufted and bipolar arrangements most common), but no basis for subdividing them into more than one class was found with any of the techniques used. The cell body and dendrites are commonly aligned parallel to the surface of TRN and at right angles to the traversing fibre bundles. The dendrites do not branch extensively and are only moderately spinous. Long, hair-like spines corresponding to those described by Scheibel & Scheibel (1966) were not found: nor were dendritic bundles found to be as prominent in EM material as reported by these authors in LM-Golgi material. Plasma membranes of dendrites in small bundles and of contiguous somata were commonly in direct contact over large areas, but gap junctions between them were not seen.The neuropil of TRN is simple with three major axon terminal types.D-type terminals (about 56% of all terminals in visual TRN) have closely packed spherical synaptic vesicles (42 nm diameter);L-type terminals (about 31%) are paler, slightly larger and have less densely packed synaptic vesicles (46 nm diameter); both terminal types make Gray type 1 synaptic contacts on dendritic spines and dendritic shafts and rarely also on cell bodies and axon hillocks.F-type terminals (about 8%) contain flattened synaptic vesicles in a dark matrix and make Gray type 2 contacts with dendrites, cell bodies and axon hillocks. In visual TRN, D-type terminals (but not all) degenerate after ablation of ipsilateral visual cortex and L-type terminals (but not all) degenerate after lesion of ipsilateral dorsal lateral geniculate nucleus; the density of degenerating terminals is higher after cortical than after geniculate lesions. Indirect evidence suggests that F-type terminals may be (or may include) collaterals of reticulo-thalamic projection cells, but no evidence was found for a widespread or dense plexus of such collaterals.After injection of HRP into the dorsal lateral geniculate nucleus, labelled axon terminals in visual TRN (many clearly L-type) were found in synaptic contact with retrogradely labelled dendrites of reticulo-geniculate projection cells. When HRP injection was combined with ablation of ipsilateral visual cortex, degenerating axon terminals (most of them identifiable as D-type) were also found in synaptic contact with retrogradely-labelled dendrites of reticulo-geniculate projection cells.Thus, neurons of visual TRN in the rat receive monosynaptic, presumptively excitatory input from collaterals of cortico-geniculate and geniculo-cortical axons, and project in a topographically-organized manner to the ipsilateral dorsal lateral geniculate nucleus (where they make Gray type 2 GABAergic and presumptively inhibitory synaptic contacts chiefly with the dendrites of geniculo-cortical projection cells). A similar pattern of organization is seen in other parts of the TRN and these data are compatible with the view that the TRN (and the perigeniculate nucleus of the cat thalamus, which is similar in several respects to visual TRN) forms part of a negative feed-back system by which the activity of thalamo-cortical projection neurons is regulated.     

Functional distinction of perigeniculate and thalamic reticular neurons in the cat

Summary Two types of neurons can be recognized in the region above the lateral geniculate nucleus. One cell type is found in the caudal part of the reticular nucleus of thalamus; these cells are accordingly called reticular neurons. The other cell type is located in the perigeniculate nucleus immediately above lamina A of the lateral geniculate nucleus and in the intermediate zone between the perigeniculate nucleus and the reticular nucleus. These cells are referred to as perigeniculate neurons. Electrical stimulation of the optic tract and the visual cortex typically evokes a short burst of spikes in the perigeniculate neurons, and the excitation has a shorter latency from the cortex (range 1.2–2.5 ms) than from the optic tract (range 1.5–3.1 ms). The perigeniculate neurons are also activated by adequate visual stimuli. In contrast, the reticular neurons are unresponsive to visual stimuli and electrical stimulation of the optic tract but they may respond with a burst of spikes to cortex stimulation with rather long latency (range 2.7–5.5 ms). It is concluded that only perigeniculate neurons qualify as interneurons in the recurrent inhibitory pathway to principal cells in the lateral geniculate nucleus.Supported by Magnus Bergvalls Stiftelse and The Swedish Medical Research Council (Project no. 4767)F.-S. Lo had an exchange fellowship from the Royal Swedish Academy of Engineering     

A study of the organization of the locus coeruleus projections to the lateral geniculate nuclei in the albino rat

The lateral geniculate nuclei of the rat are known to receive an innervation from catecholamine-containing neurons. In the present study the origin, axonal projections and terminal distribution of this innervation was studied. The lateral geniculate nuclei contain a356 ± 20 ng norepinephrine/g and64 ± 7 ng dopamine/g tissue; the latter is within the range expected for dopamine as a precursor in a region innervated by a norepinephrine-containing terminal system. When separate norepinephrine-containing cell groups located at various brain stem levels are ablated or their axonal projections destroyed, only lesions in the locus coeruleus produce a significant decrease in the norepinephrine content of the lateral geniculate nuclei. Injections of horseradish peroxidase into the lateral geniculate nuclei result in retrograde transport of horseradish peroxidase only to the noradrenergic neurons of the locus coeruleus. The labelled neurons are pretent throughout the rostrocaudal and dorsoventral axes of both the ipsilateral (60%) and contralateral (40%) nucleus. Autoradiographic and fluorescence histo-chemical experiments indicate that axons that ascend from the locus coeruleus reach the lateral geniculate nuclei via the dorsal tegmental catecholamine-containing bundle and the medial forebrain bundle. These fibers enter the ventral lateral geniculate nucleus from the zona incerta and the dorsal lateral geniculate nucleus from the superior thalamic radiation, thalamic reticular nucleus, and lateral posterior nucleus. Contralateral fibers from the locus coeruleus cross in the posterior commissure, supraoptic and pontine decussations and join the ipsilateral projections to the lateral geniculate nuclei. The bilateral locus coeruleus innervation of the nuclei is comprised of a highly branched network of varicose axons. Neither the ipsilateral nor the contralateral projections appear to be topographically organized; instead, a single fiber may have collateral axons that branch throughout large areas of the nuclei. This innervation is moderately dense in the ventral, and very dense in the dorsal, lateral geniculate nucleus.The study indicates that both the dorsal and ventral lateral geniculate nuclei receive a diffuse catecholamine-containing innervation which arises solely from the norepinephrine-containing neurons of the locus coeruleus. The innervation of each lateral geniculate nucleus is bilateral, with noradrenergic neurons located throughout both the ipsilateral and the contralateral locus coeruleus contributing to ascending pathways that terminate as a diffuse, plexiform innervation interspersed among other afferents to the lateral geniculate nuclei. It is speculated that such a diffuse noradrenergic innervation might depress the spontaneous activity of neurons in the lateral geniculate nuclei, while preserving or enhancing their responsiveness to afferent optic stimulation.     

The organization of the crossed geniculogeniculate pathway of the rat: a Phaseolus vulgaris-leucoagglutinin study.

The intergeniculate leaflet of the thalamus is known to give rise to neuronal projections to the suprachiasmatic nuclei and the rostral part of the pineal gland. Via these projections the intergeniculate leaflet is considered to play a role in regulation of circadian rhythms. Iontophoretic injections of the anterograde tracer Phaseolus vulgaris-leucoagglutinin were placed in various subnuclei of the lateral geniculate nucleus in order to study the topographical organization of the crossed geniculogeniculate pathway in the rat. Injections involving neurons in the intergeniculate leaflet or the medial subpart of the ventral nucleus (which presumably is part of the intergeniculate leaflet of the thalamus too) gave rise to labeled nerve fibers in the opposite lateral geniculate nucleus. The axons contained in this pathway were followed either medially via the posterior commissure, or via the optic tracts and optic chiasm, to the contralateral hemisphere. In the contralateral lateral geniculate nucleus, the intergeniculate leaflet was most densely innervated, but a substantial innervation of the ventral lateral geniculate nucleus was observed as well. Only a few labeled fibers were observed in the dorsal subnucleus. However, the dense innervation of the contralateral intergeniculate leaflet not only covered the small zone between the dorsal and ventral nuclei, but also a dorsomedial part of the ventral nucleus that merged caudally with the lateral part of the zona incerta. In the remaining part of the ventral nucleus, single Phaseolus vulgaris-leucoagglutinin-labeled fibers surrounded specific cells. The demonstration of a divergent projection between the intergeniculate leaflet and specific subparts of the contralateral geniculate nuclei indicates that the two lateral geniculate nuclei are regulating each other. The function of this pathway is suggested to be related to the regulation of circadian rhythmicity, but experimental evidence for this hypothesis is still lacking.     

Comparison of cholinergic and histaminergic axons in the lateral geniculate complex of the macaque monkey.

The cholinergic and histaminergic projections have important neuromodulatory functions in the ascending visual pathways, so we compared the pattern and mode of innervation of the two projections in the lateral geniculate complex (dorsal lateral geniculate nucleus and pregeniculate nucleus) of the macaque monkey. Brain tissue from macaques was immunoreacted by means of antibodies to choline acetyltransferase (ChAT) or to histamine and processed for light and electron microscopy. A dense plexus of thin, highly branched ChAT-immunoreactive axons laden with varicosities was found in all layers of the dLGN including the koniocellular laminae and in the pregeniculate nucleus. ChAT label was more dense in magnocellular layers 1 and 2 than in parvocellular layers 3-6 and relatively sparse in the interlaminar zones. Varicosities associated with the cholinergic axons had an average of three conventional asymmetric synapses per varicosity, and these appeared to contact dendrites of both thalamocortical cells and interneurons. Histamine-immunoreactive axons were distributed homogeneously throughout all laminar and interlaminar zones of the dLGN, but were denser in the pregeniculate nucleus than in the dLGN. Histaminergic axons branched infrequently and were typically larger in caliber than cholinergic axons. The overwhelming majority of varicosities were found en passant and rarely displayed conventional synapses, despite the abundance of synaptic vesicles, and were not associated preferentially with specific cellular structures. The innervation of the macaque dLGN complex by cholinergic and histaminergic systems is consistent with their proposed role in state dependent modulation of thalamic activity. The dense and highly synaptic innervation by cholinergic axons supports the proposal of additional involvement of these axons in functions related to eye movements.