PubMed Health⌕ Search

Biomedical subjects

H J Karten

Publications and source records attributed to H J Karten.

At least 19 recordsLinked to original sources
PubMed2003

Tracing developing pathways in the brain: a comparison of carbocyanine dyes and cholera toxin b subunit.

The present study examined the efficiency of fluorescent carbocyanine dye 1,1'-dioctadecyl-3,3,3',3'-tetramethylinodocarbocyanine perchlorate and cholera toxin B subunit in tracing the crossed tectal projection to the nucleus rotundus of the thalamus (tectorotundal pathways) of paraformaldehyde-fixed and living chick embryos. The tracers were injected into the optic tectum under three experimental conditions (carbocyanine postfix, carbocyanine in vivo, and cholera toxin B subunit in vivo) and the anterograde transport of the nucleus rotundus was monitored and compared. In the carbocyanine postfix method, small crystals of carbocyanine dye were inserted into the tectum of paraformaldehyde-fixed embryos. A 6-month post-insertion period was required to label the crossed tectorotundal pathway. Results showed that tectal neurons did not begin to innervate the ipsilateral nucleus rotundus until embryonic day 9 and the contralateral nucleus rotundus until embryonic day 17. This slow progression of labeling through the crossed tectal projection resulted in significant contrast of the labeling between the ipsilateral and contralateral nuclei rotundus. In the carbocyanine in vivo method, a small volume of carbocyanine dye solution was injected into the tectum of living embryos. A 8- to 12-h survival period was sufficient enough to label the tectorotundal pathway. By embryonic day 8, the labeled axons terminated in the ipsilateral nucleus rotundus and the crossed tectorotundal projection was first detected by embryonic day 10. Similarly, in the cholera toxin B subunit in vivo method, a small volume of cholera toxin B subunit solution was injected into the tectum of living embryos. After a 6- to 10-h survival period, heavily labeled axons were found to innervate bilaterally the nucleus rotundus by embryonic day 8. This appeared to be the earliest schedule for detecting the crossed tectorotundal projection, compared with that of both the postfix and in vivo methods of carbocyanine dye. Based on the differences in the detectability of the crossed tectorotundal projection between the postfix and in vivo methods, the present data suggest that the former method is of limited purpose for labeling tectal collaterals during embryogenesis. Moreover, given the rapid transport rate and absence of photobleaching, which is often seen when using carbocyanine dye, the cholera toxin B subunit in vivo method appears to be the tracer of choice for investigating embryonic pathways.

Animals↗
PubMed2001

Chattering and differential signal processing in identified motion-sensitive neurons of parallel visual pathways in the chick tectum.

At least three identified cell types in the stratum griseum centrale (SGC) of the chick optic tectum mediate separate pathways from the retina to different subdivisions of the thalamic nucleus rotundus. Two of these, SGC type I and type II, constitute the major direct inputs to rotundal subdivisions that process various aspects of visual information, e.g., motion and luminance changes. Here, we examined the responses of these cell types to somatic current injection and synaptic input. We used a brain slice preparation of the chick tectum and applied whole-cell patch recordings, restricted electrical stimulation of dendritic endings, and subsequent labeling with biocytin. Type I neurons responded with regular sequences of bursts ("chattering") to depolarizing current injection. Electrical stimulation of retinal afferents evoked a sharp-onset EPSP/burst response that was blocked with CNQX. The sharp-onset EPSP/burst response to synaptic stimulation persisted when the soma was hyperpolarized, thus suggesting the presence of dendritic spike generation. In contrast, the type II neurons responded to depolarizing current injection solely with an irregular sequence of individual spikes. Electrical stimulation of retinal afferents led to slow and long-lasting EPSPs that gave rise to one or several action potentials. In conclusion, the morphological distinct SGC type I and II neurons also have different response properties to retinal inputs. This difference is likely to have functional significance for the differential processing of visual information in the separate pathways from the retina to different subdivisions of the thalamic nucleus rotundus.

Action Potentials↗
PubMed2001

Nerve growth factor induces light adaptive cellular and synaptic plasticity in the outer retina of fish.

Recent evidence suggests that neurotrophins can be involved in short-term synaptic plasticity in parts of the central nervous system. In the present study, the possible role of nerve growth factor (NGF) in inducing morphologic (cellular and subcellular) changes in the outer retina of carp was assessed. The effects of NGF on cone photomechanical movements (PMMs) and horizontal cell (HC) spinule formation were measured. NGF-induced cone contraction and formation of HC spinules in the dark-adapted retina were consistent with its role in light adaptation. These effects were dose dependent in the range of 5--250 nM. Because cone contraction and HC spinule formation have previously been shown to be controlled by dopamine (DA), nitric oxide (NO), or both, the possibility that the effects of NGF could be occurring by means of release of DA and/or NO was tested. Haloperidol (HAL), a nonspecific DA receptor blocker, or 2-(4-carboxyphenyl)-4,4,5,5-tetramethyl-imidazoline-1-oxyl-3-oxide potassium (cPTIO), a NO scavenger, was applied in combination with NGF to dark-adapted eyecups. The results showed that both HAL and cPTIO significantly blocked the effects of NGF on cone PMMs and HC spinule formation. In conclusion, (1) NGF represents a novel light-adaptive signalling mechanism in the outer retina of fish; and (2) NGF-induced cone contraction and HC spinule formation in the retina together with our previous observation would suggest that the effects of NGF may be mediated through NO by means of DA.

Adaptation, Ocular↗
PubMed2000

Bottlebrush dendritic endings and large dendritic fields: motion-detecting neurons in the mammalian tectum.

The widefield vertical neurons of the lower stratum griseum superficiale (SGS3) and upper stratum opticum (SO) of the superior colliculus provide an extrageniculate route for visual information to reach the pulvinar. Previous physiological studies indicate that SGS3/SO neurons have large receptive fields and respond to small moving stimuli. We sought to better characterize the dendritic morphology of SGS3/SO neurons with intracellular filling in slice preparations of the ground squirrel superior colliculus. We found that dendrites of widefield vertical cells end in monostratified arrays of spiny terminal specializations called "bottlebrush" dendritic endings. Two major subtypes of neurons are described. Type I neurons have somata restricted to the SGS3 and bottlebrush endings in the most superficial sublayer of the SGS. Type II neurons are found at the base of the SGS and in the upper SO, and have bottlebrush endings arrayed within the middle sublayers of the SGS. Bottlebrush endings may sample and integrate laminated afferents to the superior colliculus, and cellular subtypes may underlie multiple information streams within the tectopulvinar pathway. A similar dendritic morphology and projection pattern can be found in cells of the avian optic tectum that project upon the nucleus rotundus, a thalamic nucleus homologous to the mammalian caudal/inferior pulvinar. Because motion processing is a dominant feature of the avian tectorotundal pathway, the current results suggest that both dendritic morphology and motion processing are conserved features of widefield vertical cells in the tectopulvinar pathway of vertebrates.

Animals↗
PubMed2000

Ontogeny of the tectorotundal pathway in chicks (Gallus gallus): birthdating and pathway tracing study.

The avian tectorotundal system has been suggested as a homologue of the mammalian colliculopulvinar system. In the tectorotundal system, neurons of the stratum griseum centrale (SGC) of the optic tectum send their axons bilaterally to the nucleus rotundus (Rt). In transit to the Rt, the axons of the SGC neurons collateralize in the nuclei posteroventralis thalami (PV), subpretectalis (SP), and interstitiopretectosubpretectalis (IPS) of the tectothalamic tract (TT). The current study used birthdating and pathway-tracing methods to investigate the neurogenesis and time course of neuronal connections of the tectorotundal pathway in chicks during embryogenesis. By using tritiated thymidine autoradiography, we observed that the SGC neurons of the tectum were generated by embryonic days 3.0-5.5 (E3.0-E5.5), the Rt by E3.5-E5.0, and the nuclei of TT by E3.5-4.5. To trace the tectorotundal pathway, we injected cholera toxin B subunit (CTb) into the tectum, and the CTb-like immunoreactivity was examined. By E4.5-E5.5, some CTb-like immunoreactive (CTb-LI) axons terminated in the ipsilateral SP/IPS. By E6.0-E6.5, CTb-LI axon bundles were seen ipsilaterally in the TT. Increased numbers of labeled axons were seen terminating in the SP/IPS. By E7.0-E7.5, heavily labeled axons in the TT were observed with diffuse terminals in areas ventral to the presumptive Rt and PV. By E7.5-E8.0, the tectal axons innervated the ipsilateral Rt, in which some of the collaterals crossed the midline to the contralateral diencephalon. The crossed tectorotundal projection was seen first by E8.0-E8.5. Also, during this stage, a few CTb-LI collaterals terminated in the contralateral SP/IPS. Between E10 and E13, the pattern of bilateral tectorotundal projections became more regionalized, whereas labeling continued to increase in the SP/IPS. At E16, the labeling pattern of all tectorecipient structures resembled that of the hatchling. The current study revealed the temporal order of development of the tectorotundal pathway during embryogenesis. The SGC cells first innervate ipsilaterally the SP/IPS and then the Rt/PV. The schedule of the crossed tectorotundal connections coincides with the schedule of tectal projections onto the contralateral intrinsic nuclei of the TT. We conclude that E8.0 (+/- E0.5) is a critical stage for the development of the tectofugal pathway. Moreover, the current study provides important insights into the relative ontogeny of the mammalian tectofugal pathway.

Animals↗
PubMed1999

Cannabinoid CB1 receptors and ligands in vertebrate retina: localization and function of an endogenous signaling system.

CB1, a cannabinoid receptor enriched in neuronal tissue, was found in high concentration in retinas of rhesus monkey, mouse, rat, chick, goldfish, and tiger salamander by using a subtype-specific polyclonal antibody. Immunolabeling was detected in the two synaptic layers of the retina, the inner and outer plexiform layers, of all six species examined. In the outer plexiform layer, CB1 was located in and/or on cone pedicles and rod spherules. Labeling was detected in some amacrine cells of all species and in the ganglion cells and ganglion cell axons of all species except fish. In addition, sparse labeling was found in the inner and/or outer segments of the photoreceptors of monkey, mouse, rat, and chick. Using GC/MS to detect possible endogenous cannabinoids, we found 3 nmol of 2-arachidonylglycerol per g of tissue, but no anandamide was detectable. Cannabinoid receptor agonists induced a dramatic reduction in the amplitude of voltage-gated L-type calcium channel currents in identified retinal bipolar cells. The presence and distribution of the CB1 receptor, the large amounts of 2-arachidonylglycerol found, and the effects of cannabinoids on calcium channel activity in bipolar cells suggest a substantive role for an endogenous cannabinoid signaling system in retinal physiology, and perhaps vision in general.

Animals↗
PubMed1999

The transport rate of cholera toxin B subunit in the retinofugal pathways of the chick.

This study investigated the transport rate of the tracer, cholera toxin B subunit, within the retinofugal pathway of the chick hatchlings. Following intraocular injections, the chicks were allowed to survive for various time-periods. The immunoreactivity of cholera toxin B subunit was then examined in the retinofugal pathways. Two hours post-injection, retinal ganglion cells began to take up the tracer and transport it to the most rostroventral portion of the optic tectum. After a 4 h survival period, the labeled retinal axons progressively innervated all retinofugal targets. Within the tectum, the labeling density varied from layer to layer with heavily labeled terminals in layer 5b, less label in layer 7 and the most diffuse label in layers 2-4. Scattered labeling was seen in the nucleus dorsolateralis anterior thalami, pars lateralis, the nucleus geniculatus lateralis, pars ventralis, the nucleus basal optic root, the nucleus lateralis anterior thalami, and the pretecal lentiformis nucleus of mesencephalon. After 6- and 8 h survival periods, increased labeling was seen in all retinofugal nuclei. There were increased numbers of retinal terminals in all retinorecipient layers of the tectum. It was noted that some of the retinal axons "overshot" into layers deeper than layer 7. In addition, retinal projections were found scattered throughout the ipsilateral nucleus basal optic root. Maximal labeling in all retinofugal targets was observed at a 10 h survival period. The present study suggests that cholera toxin B subunit can be used to trace retinal axons along their retinofugal paths up to the small terminal zones at a rate of 4.25 mm/h or 102 mm/day. Also, evidence of synchronous retinal terminations in layers 5b and 7 indicates that the transport of cholera toxin B subunit is independent of axon diameters of retinal ganglion cells. Finally, given the changing status of the embryo, the rapid transport of cholera toxin B subunit can be applied for tracing developing pathways.

Animals↗
PubMed1998

Nerve growth factor receptor TrkA is expressed by horizontal and amacrine cells during chicken retinal development.

Nerve growth factor is known to stimulate neurite outgrowth and support neuronal survival during embryonic development. We have studied the expression of the nerve growth factor receptor, TrkA, at both mRNA and protein levels during the course of chicken retinal development. Furthermore, we have compared the expression of trkA mRNA with that of the 75-kD low-affinity neurotrophin receptor (p75NTR). RNase protection assay identified peak-levels of trkA mRNA in the late embryonic retina. Using in situ hybridization and immunohistochemistry, we found cells expressing TrkA in both the internal and the external part of the inner nuclear layer, corresponding to amacrine and horizontal cells, respectively. The TrkA-expressing amacrine cell has a unistratified dendritic arborization in the second sublamina of the inner plexiform layer, and may represent the stellate amacrine cell described by Cajal. The horizontal cells, possessing arciform dendrite processes in the outer plexiform layer, showed strong TrkA immunoreactivity in both dendrites and cell bodies. During the course of retinal development, the TrkA-expressing amacrine cells decreased in number, whereas the TrkA-expressing horizontal cells persisted. Because nerve growth factor was expressed where the horizontal cells, but not where the amacrine cells were located, these findings raise the question of whether nerve growth factor could locally support the survival of TrkA-expressing interneurons during retinal development.

Animals↗
PubMed1998

Bottlebrush dendritic endings and large dendritic fields: motion-detecting neurons in the tectofugal pathway.

In avian and mammalian brains, visual information from the retina is conveyed to the telencephalon via two separate pathways: the thalamofugal and the tectofugal pathways. Recently, Karten et al. ([1997] J. Comp. Neurol. 387:449-465) examined a portion of the tectofugal pathway, the projection from the optic tectum to the nucleus rotundus thalami, in pigeons. They defined two distinct subpopulations of tectal neurons projecting from the stratum griseum centrale (SGC; tectal layer 13) to specific divisions of the rotundus. The goal of this study in chick was to verify the existence of the type I and type II SGC neurons, as defined by Karten et al., and then examine in greater detail the connectivity and morphology of these SGC neurons. Furthermore, our results suggest how the unique morphological characteristics of SGC neurons contribute to the large receptive fields (20-50 degrees) found in physiological recordings and the SGC neuronal response to extremely small (ca. 0.05 degree), fast-moving (100 degrees/second) stimuli. By injecting retrograde tracer into various divisions of the chick rotundus, we verified that, indeed, the chick did possess type I and type II SGC neurons, as well as a "new" type of SGC neuron, type III, that is not found in the pigeon. We then used intracellular cell-filling techniques to define further these three types of SGC neurons. Our examination revealed the following: Type I SGC neurons had large, circular dendritic fields (average diameter, 1,725 microns) composed of smooth dendrites and ending in spine-rich, bottlebrush endings located in retinorecipient tectal layer 5b; type II SGC neurons had elliptical dendritic fields (average 1,447 microns) and dendritic endings located never more superficially than tectal layer 8; and type III SGC neurons had large dendritic fields (average 1,800 microns) of unknown shape and bottlebrush dendritic endings located in retinorecipient tectal layer 4. We suggest that the neuronal features of the SGC neurons (i.e., bottlebrush dendritic endings and large dendritic fields) are key morphological characteristics for the detection of motion within the tectofugal pathway. Furthermore, because neurons with similar morphology have also been found in the tecta of both mammals and reptiles, we suggest that these neuronal features are fundamental components of a phylogenetically conserved system used for the "extrastriate" detection of motion in vertebrates.

Animals↗
PubMed1998

Distribution of the alpha7 nicotinic acetylcholine receptor subunit in the developing chick cerebellum.

Previous studies of the nicotinic acetylcholine receptor (nAChR) subunits in adult mammalian and avian brains have demonstrated a spatially restricted distribution of these subunits; little, however, is known about the nAChR subunit developmental distribution. The present study demonstrated a transient pattern of distribution of the neuronal nAChR subunit, alpha7, in the developing chick cerebellum by using immunohistochemical techniques. This transient distribution may suggest a critical period for the development of the cholinergic system in the cerebellum.

Animals↗
PubMed1997

Differential co-localization of nicotinic acetylcholine receptor subunits with calcium-binding proteins in retinal ganglion cells.

Immunohistochemistry was used to examine the co-occurrence of nicotinic acetylcholine receptor subunits with calcium-binding proteins in ganglion cells of the chick retina. The alpha3 subunit was rarely observed in ganglion cells containing calbindin, calretinin, or parvalbumin. On the other hand, the alpha8 subunit was more often co-localized with all calcium-binding proteins studied. These results may be related to the high calcium permeability of nicotinic receptors that contain the alpha8 subunit.

Animals↗
PubMed1997

Two distinct populations of tectal neurons have unique connections within the retinotectorotundal pathway of the pigeon (Columba livia).

The tectofugal pathway is a massive ascending polysynaptic pathway from the tectum to the thalamus and then to the telencephalon. In birds, the initial component of this pathway is known as the tectorotundal pathway; in mammals, it is known as the tectopulvinar pathway. The avian tectorotundal pathway is highly developed; thus, it provides a particularly appropriate model for exploring the fundamental properties of this system in all amniotes. To further define the connectivity of the tectorotundal projections of the tectofugal pathway, we injected cholera toxin B fragment into various rotundal divisions, the tectobulbar projection, and the ventral supraoptic decussation of the pigeon. We found intense bilateral retrograde labeling of neurons that stratified within layer 13 and, in certain cases, granular staining in layer 5b of the optic tectum. Based on these results, we propose that there are two distinct types of layer 13 neurons that project to the rotundus: 1) type I neurons, which are found in the outer sublamina of layer 13 (closer to layer 12) and which project to the anterior and centralis rotundal divisions, and 2) type II neurons, which are found in the inner sublamina of layer 13 (closer to layer 14) and which project to the posterior and triangularis rotundal divisions. Only the labeling of type I neurons produced the granular dendritic staining in layer 5b. An additional type of tectal neuron was also found that projected to the tectobulbar system. We then injected Phaseolus vulgaris-leucoagglutinin in the optic tract and found that the retinal axons terminating within tectal layer 5b formed narrow radial arbors (7-10 microm in diameter) that were confined to layer 5b. Based on these results, we propose that these axons are derived from a population of small retinal ganglion cells (4.5-6.0 microm in diameter) that terminate on the distal dendrites of type I neurons. This study strongly indicated the presence of a major bilateral oligosynaptic retinotectorotundal pathway arising from small retinal ganglion cells projecting to the rotundus with only a single intervening tectal neuron, the proposed type I neuron. We suggest that a similar organization of retinotectopulvinar connections exist in reptiles and in many mammals.

Afferent Pathways↗
PubMed1996

GABAergic inputs to the nucleus rotundus (pulvinar inferior) of the pigeon (columba livia).

The avian nucleus rotundus, a nucleus that appears to be homologous to the inferior/ caudal pulvinar of mammals, is the major target of an ascending retino-tecto-thalamic pathway. Further clarification of the inputs to the rotundus and their functional properties will contribute to our understanding of the fundamental role of the ascending tectal inputs to the telencephalon in all vertebrates, including mammals. We found that the rotundus contains a massive plexus of glutamic acid decarboxylase (GAD)-immunoreactive axons using antibodies against GAD. The cells within the rotundus, however, were not immunoreactive for GAD. The retrograde tracer cholera toxin B fragment was injected into the rotundus to establish the location of the afferent neurons and determine the source of the gamma-aminobutyric acid (GABA) inputs into the rotundus. In addition to the recognized bilateral inputs from layer 13 of the tectum, we found intense retrograde labeling of neurons within the ipsilateral nuclei subpretectalis (SP), subpretectalis-caudalis (SPcd), interstitio-pretecto-subpretectalis (IPS), posteroventralis thalami (PV), and reticularis superior thalami (RS). All the neurons of the SP, SPcd, IPS, and PV were intensely GAD-immunoreactive. The neurons of layer 13 of the tectum were not immunoreactive for GAD. Following the destruction of the ipsilateral SP/IPS complex, we found a major reduction in the intensity of the GAD axonal immunoreactivity within the ipsilateral rotundus, but this destruction did not diminish the intensity of the GAD-immunoreactivity within the contralateral rotundus. Our studies indicated that the source of the massive GAD-immunoreactive plexus within the rotundus was from the ipsilateral SP, SPcd, IPS, and PV nuclei. These nuclei, in turn, received ipsilateral tectal input via collaterals of the neurons of layer 13 in the course of their projections upon the rotundus. We suggest that the direct bilateral tecto-rotundal projections are excitatory, whereas the indirect ipsilateral projections from the SP/IPS and PV are mainly inhibitory, possibly acting via a GABA-A receptor.

Animals↗
PubMed1996

Projection of the nucleus pretectalis to a retinorecipient tectal layer in the pigeon (Columba livia).

The avian optic tectum is composed of at least 15 separate laminae that are distinguishable on the basis of their morphological features and patterns of afferent and efferent connectivity. Layer 5b, a major retinorecipient layer, exhibits dense, dust-like, neuropeptide Y-positive (NPY+) immunoreactive labeling, whereas sparse, larger caliber NPY+ fibers are found in laminae 4 and 7. Anterograde and retrograde labeling techniques, immunohistochemistry, and retinal lesion studies were used to determine the source of this tectal NPY+ labeling. NPY+ was not detectable in cells of the optic tectum or in retinal ganglion cells, and retinal ablation did not diminish the abundance of tectal NPY+ fibers. Neurons of two nuclei previously shown to be sources of tectal input, the nucleus pretectalis (PT) and the intergeniculate leaflet (IGL; Brecha, 1978), were found to be NPY+. Unilateral injection of retrograde tracers into the tectum resulted in bilateral labeling of neurons within PT, and injections of anterograde tracer into PT confirmed that this nucleus projected bilaterally to layer 5b of the optic tectum. Unilateral lesions of PT nearly eliminated NPY+ fibers in the ipsilateral layer 5b and significantly reduced them in the contralateral layer 5b. Bilateral lesions of PT eliminated NPY+ fibers bilaterally in layer 5b. However, these PT lesions had little effect on the NPY+ fibers in layers 4 and 7. Combined retrograde and immunohistochemical studies showed that NPY+ neurons of the IGL project to the optic tectum, and anterograde studies demonstrated that IGL projects to layers 4 and 7. The NPY+ projection to laminae 5b from PT is one of many inputs, which include cholinergic afferents from the nucleus isthmi parvicellularis, terminals from retinal ganglion cells, and dendrites of layer 13 neurons (Karten et al., 1993). The NPY+ input to layer 5b may modulate visual information flow from retinal input to various tectal neurons, including those in layer 13.

Animals↗
PubMed1995

Centrifugal projections upon the retina: an anterograde tracing study in the pigeon (Columba livia).

Previous work has shown that the avian retina receives two types of centrifugal fibers from the brain. These types can be distinguished based on the size and the morphology of their terminal endings and have been termed convergent and divergent. The centrifugal fibers arise from the isthmooptic nucleus (ION) and the surrounding ectopic cell region (ECR). We used injections of anterograde tracers either to the ION/ECR or to the ECR only to determine the morphology, depth of termination, and regional distribution of the centrifugal fibers arising from each. We found that the ECR gives rise only to the divergent type of the centrifugal fiber, whereas the ION gives rise mainly to the convergent type but may also send some fibers of the divergent type. Most of the fibers project contralaterally, although a few from the ECR project ipsilaterally. The terminals of either type are not uniformly distributed throughout the retina; instead, they are found mainly in the inferior, midtemporal, to nasal portion of the retina and appear to avoid the fovea and most of the red field. By comparison, the ION receives a major projection from portions of the tectum that receive input from the fovea and the red field in a type of neural loop. The neural loop does not project to the same point (homotopic), but projects from the red field to the inferior retina (heterotopic), as was recently proposed by Holden (1990; Vis. Neurosci. 4:493-497). The distribution of centrifugal axons corresponds to displaced ganglion cells that selectively innervate the nuclei of the accessory optic system (AOS), including the nucleus of the basal optic root (dorsal, ventral, and lateral) and the nucleus lentiformis mesencephali, pars magnocellularis. We suggest that the centrifugal axons act by increasing the gain on the AOS, thereby enhancing retinal stabilization of gaze with improved accuracy of pecking of small objects.

Animals↗
PubMed1995

Intratelencephalic projections of the visual wulst in pigeons (Columba livia).

The visual wulst is the telencephalic target of the thalamofugal visual pathway of birds, and thus the avian equivalent of the striate cortex of mammals. The anterograde tracer Phaseolus vulgaris leucoagglutinin was used to follow the intratelencephalic connections of the major constituents of the visual wulst in pigeons. In particular, efferent pathways from the granular layer (Intercalated nucleus of the hyperstriatum accessorium, IHA), supragranular layer (hyperstriatum accessorium, HA), and infragranular layers (hyperstriatum intercalatus superior and/or hyperstriatum dorsale, HIS/HD) were investigated. These efferent projections were confirmed by injections of the retrograde tracer cholera toxin subunit B into their terminal fields. When a deposit of the anterograde tracer was centered in IHA, which receives the visual thalamic input, efferent fibers were seen mainly dorsomedially to IHA. When a deposit of the anterograde tracer was centered in HA, efferent fibers were seen to extend mainly in three directions: 1) medially to the tractus septomesencephalicus, which sends projections to extratelencephalic visual nuclei: 2) ventrolaterally to the lateral portion of the neostriatum frontale, where there were also labeled cells after the retrograde tracer was injected in HA; and 3) ventromedially to the paleostriatal complex, which is the avian equivalent of the mammalian caudale, 5) neostriatum intermedium, 6) archistriatum intermedium, and 7) hyperstriatum laterale. Finally, HIS/HD have projections predominantly to HA and the dorsocaudal telencephalon (area corticoidea dorsolateralis and area parahippocampalis), as well as relatively minor projections to the areas which also receive projections from HA. No anterogradely labeled fibers were seen in the tractus septomesencephalicus following the tracer injections in HIS/HD. These results indicate that the visual information from the granular layer is distributed via the supragranular layer HA to multiple areas within the telencephalon, such as the neostriatum frontale and paleostriatal complex. In addition, HA is the source of an extratelencephalic projection via the tractus septomesencephalicus. Thus, the avian supragranular layer HA contains neurons which are the source of both intratelencephalic and extratelencephalic projections, whereas neurons of the mammalian cortex are segregated into two distinct layers, supragranular and infragranular layers, based on the targets of their projections. The findings are further discussed and compared to the mammalian striate cortex.

Animals↗