Biomedical subjects

Alessandra Angelucci

Publications and source records attributed to Alessandra Angelucci.

7 recordsLinked to original sources

PubMed2006

Contribution of feedforward thalamic afferents and corticogeniculate feedback to the spatial summation area of macaque V1 and LGN.

Neurons in the primary visual cortex (V1) respond best to oriented gratings of optimal size within their receptive field (RF) and are suppressed by larger gratings involving the nonclassical RF surround. A V1 neuron's optimal stimulus size is larger at lower stimulus contrast. A central question in visual neuroscience is what circuits generate the size tuning of V1 cells. We recently demonstrated that V1 horizontal connections integrate signals within a region of the RF center corresponding to the V1 neuron's optimal stimulus size at low contrast; extrastriate feedback connections to V1, instead, are longer range and can integrate signals from the most distant regions of the V1 cell's RF surround. Here, we have determined the contribution of geniculocortical feedforward and corticogeniculate feedback connections to the size-tuning of macaque V1 and lateral geniculate (LGN) neurons, respectively. Specifically, we have quantitatively compared the visuotopic extent of geniculate feedforward afferents to V1 with the size of the RF center and surround of neurons in the V1 input layers and the visuotopic extent of V1 feedback connections to the LGN with the RF size of cells in V1 layer 6, where these connections originate. We find geniculate feedforward connections to provide visuotopic information to V1 that is spatially coextensive with the V1 neuron's optimal stimulus size measured with high-contrast gratings. V1 feedback connections restrict their influence to an LGN region visuotopically coextensive with the size of the minimum response field (or classical RF) of V1 layer 6 cells and commensurate with the LGN region from which they receive feedforward connections.

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PubMed2006

The role of feedback in shaping the extra-classical receptive field of cortical neurons: a recurrent network model.

The responses of neurons in sensory cortices are affected by the spatial context within which stimuli are embedded. In the primary visual cortex (V1), orientation-selective responses to stimuli in the receptive field (RF) center are suppressed by similarly oriented stimuli in the RF surround. Surround suppression, a likely neural correlate of perceptual figure-ground segregation, is traditionally thought to be generated within V1 by long-range horizontal connections. Recently however, it has been shown that these connections are too short and too slow to mediate fast suppression from distant regions of the RF surround. We use an anatomically and physiologically constrained recurrent network model of macaque V1 to show how interareal feedback connections, which are faster and longer-range than horizontal connections, can generate "far" surround suppression. We provide a novel solution to the puzzle of how surround suppression can arise from excitatory feedback axons contacting predominantly excitatory neurons in V1. The basic mechanism involves divergent feedback connections from the far surround targeting pyramidal neurons sending monosynaptic horizontal connections to excitatory and inhibitory neurons in the RF center. One of several predictions of our model is that the "suppressive far surround" is not always suppressive, but can facilitate the response of the RF center, depending on the amount of excitatory drive to the local inhibitors. Our model provides a general mechanism of how top-down feedback signals directly contribute to generating cortical neuron responses to simple sensory stimuli.

Action Potentials↗

PubMed2006

Contribution of feedforward, lateral and feedback connections to the classical receptive field center and extra-classical receptive field surround of primate V1 neurons.

A central question in visual neuroscience is what circuits generate the responses of neurons in the primary visual cortex (V1). V1 neurons respond best to oriented stimuli of optimal size within their receptive field (RF) center. This size tuning is contrast dependent, i.e. a neuron's optimal stimulus size measured at high contrast (the high-contrast summation RF, or hsRF) is smaller than when measured using low-contrast stimuli (the low-contrast summation RF, or lsRF). Responses to stimuli in the RF center are usually suppressed by iso-oriented stimuli in the extra-classical RF surround. Iso-orientation surround suppression is fast and long range, extending well beyond the size of V1 cells' lsRF. Geniculocortical feedforward (FF), V1 lateral and extrastriate feedback (FB) connections to V1 could all contribute to generating the RF center and surround of V1 neurons. Studies on the spatio-temporal properties and functional organization of these connections can help disclose their specific contributions to the responses of V1 cells. These studies, reviewed in this chapter, have shown that FF afferents to V1 integrate signals within the hsRF of V1 cells; V1 lateral connections are commensurate with the size of the lsRF and may, thus, underlie contrast-dependent changes in spatial summation, and modulatory effects arising from the surround region closer to the RF center (the "near" surround). The spatial and temporal properties of lateral connections cannot account for the dimensions and onset latency of modulation arising from more distant regions of the surround (the "far" surround). Inter-areal FB connections to V1, instead, are commensurate with the full spatial range of center and surround responses, and show fast conduction velocity consistent with the short onset latency of modulation arising from the "far" surround. We review data showing that a subset of FB connections terminate in a patchy fashion in V1, and show modular and orientation specificity, consistent with their proposed role in orientation-specific center-surround interactions. We propose specific mechanisms by which each connection type contributes to the RF center and surround of V1 neurons, and implement these hypotheses into a recurrent network model. We show physiological data in support of the model's predictions, revealing that modulation from the "far" surround is not always suppressive, but can be facilitatory under specific stimulus conditions.

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PubMed2003

Anatomical substrates for functional columns in macaque monkey primary visual cortex.

In this review we re-examine the concept of a cortical column in macaque primary visual cortex, and consider to what extent a functionally defined column reflects any sort of anatomical entity that subdivides cortical territory. Functional studies have shown that columns relating to different response properties are mapped in cortex at different spatial scales. We suggest that these properties first emerge in mid-layer 4C through a combination of thalamic afferent inputs and local intracortical circuitry, and are then transferred to other layers in a columnar fashion, via interlaminar relays, where additional processing occurs. However, several properties are not strictly columnar since they do not appear in all cortical layers. In contrast to the functional column, an anatomically based cortical column is defined most clearly in terms of the reciprocal connections it makes, both via intra-areal lateral connections and inter-areal feedback/feedforward pathways. The column boundaries are reinforced by interplay between lateral inhibition spreading beyond the column boundary and disinhibition within the column. The anatomical column acts as a functionally tuned unit and point of information collation from laterally offset regions and feedback pathways. Thalamic inputs provide the high-contrast receptive field sizes of the column's neurons, intra-areal lateral connections provide their low contrast summation field sizes, and feedback pathways provide surround modulation of receptive fields responses.

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PubMed2002

Circuits for local and global signal integration in primary visual cortex.

Contrast-dependent changes in spatial summation and contextual modulation of primary visual cortex (V1) neuron responses to stimulation of their receptive field reveal long-distance integration of visual signals within V1, well beyond the classical receptive field (cRF) of single neurons. To identify the cortical circuits mediating these long-distance computations, we have used a combination of anatomical and physiological recording methods to determine the spatial scale and retinotopic logic of intra-areal V1 horizontal connections and inter-areal feedback connections to V1. We have then compared the spatial scales of these connectional systems to the spatial dimensions of the cRF, spatial summation field (SF), and modulatory surround field of macaque V1 neurons. We find that monosynaptic horizontal connections within area V1 are of an appropriate spatial scale to mediate interactions within the SF of V1 neurons and to underlie contrast-dependent changes in SF size. Contrary to common beliefs, these connections cannot fully account for the dimensions of the surround field. The spatial scale of feedback circuits from extrastriate cortex to V1 is, instead, commensurate with the full spatial range of center-surround interactions. Thus these connections could represent an anatomical substrate for contextual modulation and global-to-local integration of visual signals. Feedback projections connect corresponding and equal-sized regions of the visual field in striate and extrastriate cortices and cover anisotropic parts of visual space, unlike V1 horizontal connections that are isotropic in the macaque. V1 isotropic connectivity demonstrates that anisotropic horizontal connections are not necessary to generate orientation selectivity. Anisotropic feedback connections may play a role in contour completion.

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PubMed2002

Anatomical origins of the classical receptive field and modulatory surround field of single neurons in macaque visual cortical area V1.

From the analyses of our own and others' anatomical and physiological data for the macaque visual system, we arrive at a conclusion that three pathways can provide the V1 neuron with access to information from the visual field and affect its response. First, direct thalamic input can determine the size of the initial activating RF at high contrast. Second, lateral connections can enlarge the RF at low contrast by pooling information from larger regions of cortex that are otherwise ineffective when high contrast thalamic input is driving the cortical neuron. Thirdly, feedback from extrastriate cortex (possibly together with overlap or interdigitation of coactive lateral connectional fields within V1) can provide a large and stimulus specific surround modulatory field. The stimulus specificity of the interactions between the center and surround fields, may be due to the orderly, matching structure and different scales of intra-areal and feedback projection excitatory pathways. The observed activity changes of single recorded excitatory neurons could be a result of the relative weight of excitation on the excitatory neurons themselves and on local inhibitory interneurons that synapse on them. Inhibitory basket neurons, driven by the local excitatory neurons, could govern local interactions between cortical patches of different tuning properties, resulting in more distant changes in excitatory input in the laterally connected intra-areal neuronal pools.

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PubMedDate not supplied

Reaching beyond the classical receptive field of V1 neurons: horizontal or feedback axons?

It is commonly assumed that the orientation-selective surround field of neurons in primary visual cortex (V1) is due to interactions provided solely by intrinsic long-range horizontal connections. We review evidence for and against this proposition and conclude that horizontal connections are too slow and cover too little visual field to subserve all the functions of suppressive surrounds of V1 neurons in the macaque monkey. We show that the extent of visual space covered by horizontal connections corresponds to the region of low contrast summation of the receptive field center mechanism. This region encompasses the classically defined receptive field center and the proximal surround. Beyond this region, feedback connections are the most likely substrate for surround suppression. We present evidence that inactivation of higher order areas leads to a major decrease in the strength of the suppressive surround of neurons in lower order areas, supporting the hypothesis that feedback connections play a major role in center-surround interactions.

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