Intrinsic projections within visual cortex: evidence for orientation-specific local connections.

The mammalian visual cortex harbors a number of functional maps that represent distinct attributes of stimuli in the visual environment. How different functional maps are accommodated within the same cortical space, especially in species that show marked irregularities in one or more functional maps, remains poorly understood. We used optical imaging of intrinsic signals and electrophysiological techniques to investigate the organization of the maps of orientation preference, ocular dominance, and visual space in ferret. This species shows striking nonuniformity in the arrangement of ocular dominance domains and disruption of the mapping of visual space along the V1/V2 border. We asked whether these irregularities would be reflected in the organization of the map of orientation preference. The results show that orientation preference is mapped consistently within both V1 and V2, and across the interareal boundary, with no reflection of the irregularities in the other maps. These observations demonstrate the accommodation of multiple functional maps within the same cortical space without systematic geometrical relationships that necessarily constrain the organization of each representation. Furthermore, they imply that the structure of the map of orientation preference reflects the architecture and activity patterns of cortical circuits that are independent of other columnar systems established in layer 4.

Using a combination of anatomical and physiological techniques we have studied some of the neural connections subserving binocular vision in two species of artiodactyl ungulates (the sheep, Ovis sp., and the goat, Capra hircus). After monocular injections of tritiated proline, transsynaptic transport was observed bilaterally in layers 4 and 6 of visual cortical areas V1 and V2, but there were no sharply defined ocular dominance columns of the kind seen in cats and rhesus monkeys. In coronal sections there was a discontinuity in density of labelling between areas V1 and V2 corresponding to a point in the visuotopic map about azimuth –15° in the ipsilateral visual field. This discontinuity was most pronounced in the hemisphere ipsilateral to the injected eye. We conclude, therefore, that while the cortical representation of ipsilateral visual space can be explained by the retino-geniculo-cortical input pathway from the contralateral eye, the physiologically demonstrated cortical contribution to ipsilateral visual space from the ipsilateral eye cannot be explained in this way. This conclusion was reinforced by experiments using retrograde transport of horseradish peroxidase from the lateral geniculate nucleus (LGN) and medial interlaminar nucleus (MIN) to retinal ganglion cells in flattened whole mounts. These experiments revealed a sharp nasotemporal decussation in the ipsilateral retina, which could not thereby subserve any significant representation of the ipsilateral visual field. In contrast the contralateral nasotemporal decussation was smeared, with many labelled ganglion cells in the temporal retina which could subserve visual input from the ipsilateral hemifield. When we estimated the projection of the nasotemporal decussation line into visual space, we found that it was tilted from vertical by about 5°in each eye, in a similar way to that already reported in the cat. Neurophysiological recordings from binocular neurons in area V1 with different vertical eccentricities also showed that the vertical horopter (the midsagittal reference plane for binocular vision) would be tilted in life when the cyclotorsional position of the eyes was taken into account. Thus both anatomical and physiological methods concur in the prediction that ungulates have a tilted vertical horopter like that described for two other terrestrial species, the burrowing owl and the cat. Anatomical experiments reveal other similarities between the organisation of the ungulate''s visual pathways and that of the cat. For example, after tritiated proline injection in V1, we found visuotopic labelling in the claustrum, dorsal LGN, cortical area V2, and the superior colliculus. Horseradish peroxidase injections in V2 also revealed a direct input to that area from the MIN and dorsal LGN and also from layers 2, 3 and 5 of the splenial area.

Cats were reared in total darkness from birth until 4-5 mo of age (DR cats, n = 7) or with very brief visual experience (1 or 2 days) during an otherwise similar period of dark rearing [DR(1) cats, n = 3; DR(2) cats, n = 7]. Single-cell recordings were made in area 17 of visual cortex at the end of this rearing period and/or after a subsequent prolonged period of monocular deprivation. Control observations were made in normal cats (n = 3), cats reared with monocular deprivation from birth (n = 4), and cats monocularly deprived after being reared normally until 4 mo of age (n = 2). After rearing cats in total darkness, the majority of visual cortical cells were binocularly driven and the overall distribution of ocular dominance was not different from that of normal cats. Orientation-selective cells were very rare in dark-reared cats. Monocular deprivation imposed after dark rearing resulted in selective development of connections from the open eye. Most cells were responsive only to the open eye and the majority of these were orientation selective. These results were similar to, though less severe than, those found in cats reared with monocular deprivation from birth. Monocular deprivation imposed after 4 mo of normal rearing did not produce selective development of connections from the open eye in terms of either ocular dominance or orientation selectivity. In DR(1) cats visual cortical physiology was degraded in comparison to dark-reared cats after the rearing period. Most cells were binocularly driven but there was a higher frequency of unresponsive cells and a reduced frequency of orientation-selective cells. Subsequent monocular deprivation resulted in a further decrease in the number of binocularly driven cells and an increase in unresponsive cells. However, it did not produce a bias in favor of the open eye in terms of either ocular dominance or orientation selectivity. In DR(2) cats there was a high incidence of unresponsive cells and a marked loss of binocularly driven cells after the rearing period. Subsequent monocular deprivation failed to produce any significant changes.(ABSTRACT TRUNCATED AT 400 WORDS)

Noradrenaline (NA)-stimulated beta-adrenoreceptors activate adenylate cyclase via excitatory G-proteins (Gs). Activated adenylate cyclase in turn promotes the production of cAMP. Critical roles of cAMP-dependent protein kinase A (PKA) in divergent cellular functions have been shown, including memory, learning and neural plasticity. Ocular dominance plasticity (ODP) is strongly expressed in early postnatal life and usually absent in the mature visual cortex. Here, we asked whether the activation of cAMP-dependent PKA could restore ODP to the aplastic visual cortex of adult cats. Concurrent with brief monocular deprivation, each of the following cAMP-related drugs was directly and continuously infused in the adult visual cortex: cholera toxin (a Gs-protein stimulant), forskolin (a Gs-protein-independent activator of adenylate cyclase) and dibutyryl cAMP (a cAMP analogue). We found that the ocular dominance distribution became W-shaped, the proportion of binocular cells being significantly lower than that in respective controls. We concluded that the activation of cAMP cascades rapidly restores ODP to the adult visual cortex, though moderately. The finding further extends the original hypothesis that the NA-beta-adrenoreceptors system is a neurochemical mechanism of cortical plasticity.

The functional organization of the cochlear nucleus (CN) was studied with physiological recording and anatomical tracing techniques. Recordings were made from single CN neurons to examine their temporal firing patterns to tone burst stimuli and their frequency tuning characteristics. Recording loci of individual neurons were carefully monitored in order to understand how the functional properties of a cell relate to its location within the CN. We found that tonal frequencies were systematically represented in each of the three CN divisions (anteroventral, AVCN; posteroventral, PVCN; dorsal, DCN). Eight temporal response patterns were observed in CN neurons when stimulated at units' best excitatory frequencies (BF). With a few exceptions, neurons in each CN division could generate all eight firing patterns with different distributions for the three division. A focal injection of horseradish peroxidase (HRP), at the end of the physiological study, to a group of neurons possessing a similar BF in one CN division resulted in anterograde labeling of nerve terminals in the other two divisions at precisely the areas where the same frequency band was processed in these divisions. Labeled terminals in each division were closely congregated in the form of a thin slab. The slab orientation was division specific whereas its location was frequency specific, which could be predicted on the basis of physiological data. HRP injections into the DCN also resulted in retrograde labeling of somata in the AVCN and PVCN. On the other hand, only DCN neurons were retrogradely labeled when HRP was injected into the AVCN or the PVCN. These data showed how the three CN divisions are internally connected. Furthermore, retrogradely labeled cells occupied the same slabs where we found anterogradely labeled nerve terminals. Additionally, in a group of bats, HRP was injected into various functionally (i.e., BF) identified regions of the central nucleus of the inferior coliculus (IC) to clarify the type and location of CN projecting neurons. Retrogradely labeled cells in individual CN divisions likewise were arranged in slabs whose locations in the CN nuclei depended on the BFs of neurons at the injection site in the IC. These results show that slabs represent units of functional organization (i.e., tonal frequency, local connection and central projection) in the CN.

Monocular deprivation (MD) during the critical period for the development of visual cortex causes a loss of binocular response of neurons and a shift to the open eye, a normal ocular dominance (OD) shift. However, when MD is combined with chronic inactivation of the visual cortex by muscimol, the OD distribution of the neurons shifts to the deprived eye (reverse OD shift). We have previously shown that the normal OD shift is abolished by chronic infusion of the protein kinase A (PKA) inhibitor, 8-chloroadenosine-3', 5'-cyclic monophosphorothioate, Rp-isomer (Rp-8-Cl-cAMPS), into kitten visual cortex. In this study, we investigated the effect of this inhibitor on the reverse OD shift. Combination of MD and muscimol infusion into the visual cortex of 6-wk-old kittens caused a reverse OD shift that was comparable to that seen in previous studies. However, a reverse OD shift was also seen with concurrent infusion of the PKA inhibitor with muscimol. The strongest OD shift was observed in layer IV regardless of the presence or absence of the PKA inhibitor. This suggests that the dissociation of pre- and postsynaptic activities, which occurs mainly at thalamocortical synapses, induces the reverse OD shift and that inhibition of PKA does not prevent it. Presumably, an inhibition of PKA has no effect in silent cortex. We conclude that 1) an activation of PKA is not required for the induction of the reverse OD shift, and 2) the intracellular signaling mechanism underlying MD-induced OD plasticity differs between normal and reverse OD shifts.

Adaptation to Natural Binocular Disparities in Primate V1 Explained by a Generalized Energy Model

Event Abstract Back to Event The model of ocular dominance pattern formation in the presence of gradients of chemical labels. Dmitry Tsigankov1, 2* and Alexei Koulakov3 1 Bernstein Center for Computational Neuroscience, Germany 2 Max Planck Institute for Dynamics and Self-organization, Germany 3 Cold Spring Harbor Laboratory, United States The formation of ocular dominance (OD) columns in the mammalian visual cortex is thought to be driven by electrical activity of the projecting neurons. Indeed, theoretical modeling have shown that lateral cortical activity-dependent interactions are capable of producing such a periodic pattern of regions dominated by inputs from left or right eye. This scenario for OD pattern formation based on self-organization of inputs due to excitatory and inhibitory lateral interactions of Mexican-hat shape was recently confronted by several lines of experimental observations. First, the anatomical data on primary visual cortex indicate that the inhibition can have a shorter range than excitation, the regime in which the classical model fails to produce OD structure. Second, the measurements of the width of OD regions following the manipulations with the strength of the inhibitory connections are inconsistent with the predictions of the model. When the strength of inhibitory connections is increased the OD width is found to increase and when inhibition is decreased the OD width also decreases. This behavior is opposite to one predicted by the classical model. Finally, the sole role of activity-dependent self-organization in the formation of OD structure was questioned as it was observed that OD patterns can be formed in optic tectum in the presence of other factors such as gradients of interacting chemical labels.Here we present theoretical analysis of the possible interplay between genetically encoded labeling and experience-driven reorganization of the projections in the formation of OD patterns. We show that in the presence of single gradient of chemical marker the projections from two eyes are segregated into OD columns for a wide class of lateral interaction profiles. We obtain that depending on the range and strength of inhibitory and excitatory lateral connections the projecting neurons may prefer to form segregated or mixed inputs. We find the regimes when OD structure emerges for short-range inhibition and long-range excitation. We also investigate the role of lateral inhibition and excitation for different interaction profiles to find a novel regime when increase in the inhibition strength increases the width of OD columns in agreement with the experiment. Conference: Bernstein Conference on Computational Neuroscience, Frankfurt am Main, Germany, 30 Sep - 2 Oct, 2009. Presentation Type: Poster Presentation Topic: Learning and plasticity Citation: Tsigankov D and Koulakov A (2009). The model of ocular dominance pattern formation in the presence of gradients of chemical labels.. Front. Comput. Neurosci. Conference Abstract: Bernstein Conference on Computational Neuroscience. doi: 10.3389/conf.neuro.10.2009.14.146 Copyright: The abstracts in this collection have not been subject to any Frontiers peer review or checks, and are not endorsed by Frontiers. They are made available through the Frontiers publishing platform as a service to conference organizers and presenters. The copyright in the individual abstracts is owned by the author of each abstract or his/her employer unless otherwise stated. Each abstract, as well as the collection of abstracts, are published under a Creative Commons CC-BY 4.0 (attribution) licence (https://creativecommons.org/licenses/by/4.0/) and may thus be reproduced, translated, adapted and be the subject of derivative works provided the authors and Frontiers are attributed. For Frontiers’ terms and conditions please see https://www.frontiersin.org/legal/terms-and-conditions. Received: 28 Aug 2009; Published Online: 28 Aug 2009. * Correspondence: Dmitry Tsigankov, Bernstein Center for Computational Neuroscience, Göttingen, Germany, dmitry@nld.ds.mpg.de Login Required This action requires you to be registered with Frontiers and logged in. To register or login click here. Abstract Info Abstract The Authors in Frontiers Dmitry Tsigankov Alexei Koulakov Google Dmitry Tsigankov Alexei Koulakov Google Scholar Dmitry Tsigankov Alexei Koulakov PubMed Dmitry Tsigankov Alexei Koulakov Related Article in Frontiers Google Scholar PubMed Abstract Close Back to top Javascript is disabled. Please enable Javascript in your browser settings in order to see all the content on this page.

Kittens were given intramuscular injections of the N-methyl--aspartate (NMDA) antagonist MK-801 twice daily (morning and midday) during the peak of the period of susceptibility for ocular dominance changes. They were then exposed to light with one eye closed for 4 h after each injection. The ocular dominance of these kittens was shifted significantly less than that of kittens injected with saline and exposed to light over the same period at the same age. After recording a sample of cells for an ocular dominance histogram, the kittens were injected with the same dose of MK-801 that was used during rearing to observe its effect on the activity of single cells in the visual cortex. In the majority of cells (7/13) there was no significant change in activity. Positive evidence for a reduction in activity was seen in only a minority (3/13) of cells. In a separate series of experiments, dose-response curves were measured for cells in the visual cortex in response to iontophoresis of NMDA or alpha-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid (AMPA), and the effect of an injection of MK-801 on these curves was measured. MK-801, at doses similar to those used in the ocular dominance experiments, had a significant effect on the dose-response curves for NMDA, but little effect on the dose-response curves for AMPA, or the visual responses of the cells. We conclude that ocular dominance shifts can be reduced significantly by a treatment that has little effect on the level of activity of cells in the visual cortex but does specifically affect the responses of the cells to NMDA as opposed to the responses to AMPA.

Neurons in primary visual cortex are selective for stimulus orientation, and a neuron’s preferred orientation changes little when the stimulus is switched from one eye to the other. It has recently been shown that monocular orientation preferences are uncorrelated before eye opening; how, then, do they become aligned during visual experience? We aimed to provide a model for this acquired congruence. Our model, which simulates the cat’s visual system, comprises multiple on-centre and off-centre channels from both eyes converging onto neurons in primary visual cortex; development proceeds in two phases via Hebbian plasticity in the geniculocortical synapse. First, cortical drive comes from waves of activity drifting across each retina. The result is orientation tuning that differs between the two eyes. The second phase begins with eye opening: at each visual field location, on-centre cortical inputs from one eye can cancel off-centre inputs from the other eye. Synaptic plasticity reduces the destructive interference by up-regulating inputs from one eye at the expense of its fellow, resulting in binocular congruence of orientation tuning. We also show that orthogonal orientation preferences at the end of the first phase result in ocular dominance, suggesting that ocular dominance is a by-product of binocular congruence.

Heterotopic and homotopic callosal connections in rat visual cortex

Anatomical and electrophysiological studies on the reciprocal projections between the subthalamic nucleus and nucleus tegmenti pedunculopontinus in the rat

We studied how object speed is reconstructed from the responses of motion-selective cells for the generation of a behavior that is tightly linked to the speed of visual motion. In theory, the speed of an object could be estimated either from the speed tuning of the active population of motion-selective cells or from the rate of displacement of activation across the cortical map of visual space. We measured the pursuit eye movements evoked by stimuli containing two conflicting motion components: a local component designed to excite motion-selective cells with a particular speed tuning and a displacement component designed to excite cells with a sequence of spatial receptive fields. Pursuit eye movements were driven primarily by the local-motion component and were affected to only a small degree by the rate of target displacement across visual space. Extracellular single-unit recordings using the same stimuli revealed that the responses of cells in the middle temporal visual area (MT) depended primarily on the local-motion component but were influenced by the displacement component to the same degree as were pursuit eye movements. We conclude that the initiation of pursuit is consistent with a reconstruction of target speed based on the speed tuning of the active population of MT cells.

In addition to a topographic map of the retina, mammalian visual cortex contains superimposed, orderly periodic maps of features such as orientation, eye dominance, direction of motion and spatial frequency. There is evidence that these maps are overlaid so as to ensure that all combinations of the different parameters are represented as uniformly as possible across visual space. However, it is unknown to what extent geometrical factors limit the number of periodic maps which might simultaneously be present, given this constraint. This paper attempts to investigate the question by using a dimension reduction model to generate maps of simple, many- dimensional feature spaces onto a model two-dimensional cortex. The feature space included a model retina, plus N binary variables, corresponding to parameters such as ocular dominance or spatial frequency. The results suggest that geometrical factors do not sharply limit the ability of the cortex to represent combinations of parameters in spatially superimposed maps of similar periodicity. Considerations of uniform coverage suggest an upper limit of six or seven maps. A higher limit, of about nine or ten, may be imposed by the numbers of neurons (or minicolumns) available to represent each of 2(N) features within a given small region of cortex.

Sixty lesions were placed in the pulvinar nucleus or the lateral geniculate nucleus of the tree shrew, and degenerated axons and their terminals were traced to the cortex. In all but two of the animals the electrode approached the thalamic target through the cerebellum and midbrain, thus sparing the cortex and thalamic nuclei other than the target nuclei. The results show that the pulvinar nucleus projects in a topographic fashion to an extensive cortical zone that includes areas 18, 19 and several temporal areas below 19. While there are several architectonic areas in its cortical target, no corresponding subdivisions could be established within the pulvinar nucleus, and its projections are best explained in terms of a single topographic organization. Further, since the pulvinar nucleus receives fibers from the superficial layers of the superior colliculus, its extensive cortical target can be regarded as primary visual cortex. The lateral geniculate nucleus projects in a precisely topographic fashion to area 17. Further, our evidence suggests that each layer of the lateral geniculate nucleus projects to a horizontal strip within the fourth or granular layer of area striata. With respect to the question of the relation between ocular dominance and location in area 17, the pattern of projections from the lateral geniculate nucleus does not reveal ocular dominance columns in layer IV of the tree shrew. In contrast to our results Hubel and Wiesel ('68, '69) have shown that in the monkey there are vertical columns of cells in layer IV driven by either the homolateral or the contralateral eye. The case for the cat (Hubel and Wiesel, '62, '65) may be different from either of the other two species, thus posing the problem of discovering the mammalian plan and departures from the prototypical organization.