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The functional geometry of local and horizontal connections in a model of V1.

A mathematical model of interacting hypercolumns in primary visual cortex (V1) is presented that incorporates details concerning the geometry of local and long-range horizontal connections. Each hypercolumn is modeled as a network of interacting excitatory and inhibitory neural populations with orientation and spatial frequency preferences organized around a pair of pinwheels. The pinwheels are arranged on a planar lattice, reflecting the crystalline-like structure of cortex. Local interactions within a hypercolumn generate orientation and spatial frequency tuning curves, which are modulated by horizontal connections between different hypercolumns on the lattice. The symmetry properties of the local and long-range connections play an important role in determining the types of spontaneous activity patterns that can arise in cortex.

Animals↗

The neurogeometry of pinwheels as a sub-Riemannian contact structure.

We present a geometrical model of the functional architecture of the primary visual cortex (V1) and, more precisely, of its pinwheel structure. The problem is to understand from within how the internal "imminent" geometry of the visual cortex can produce the "transcendent" geometry of the external space. We use first the concept of blowing up to model V1 as a discrete approximation of a continuous fibration pi: R x P --> P with base space the space of the retina R and fiber the projective line P of the orientations of the plane. The core of the paper consists first in showing that the horizontal cortico-cortical connections of V1 implement what the geometers call the contact structure of the fibration pi, and secondly in introducing an integrability condition and the integral curves associated with it. The paper develops then three applications: (i) to Field's, Hayes', and Hess' psychophysical concept of association field, (ii) to a variational model of curved modal illusory contours (in the spirit of previous models due to Ullman, Horn, and Mumford), (iii) to Ermentrout's, Cowan's, Bressloff's, Golubitsky's models of visual hallucinations.

Animals↗

Figure-ground activity in V1 and guidance of saccadic eye movements.

Every day we shift our gaze about 150.000 times mostly without noticing it. The direction of these gaze shifts are not random but directed by sensory information and internal factors. After each movement the eyes hold still for a brief moment so that visual information at the center of our gaze can be processed in detail. This means that visual information at the saccade target location is sufficient to accurately guide the gaze shift but yet is not sufficiently processed to be fully perceived. In this paper I will discuss the possible role of activity in the primary visual cortex (V1), in particular figure-ground activity, in oculo-motor behavior. Figure-ground activity occurs during the late response period of V1 neurons and correlates with perception. The strength of figure-ground responses predicts the direction and moment of saccadic eye movements. The superior colliculus, a gaze control center that integrates visual and motor signals, receives direct anatomical connections from V1. These projections may convey the perceptual information that is required for appropriate gaze shifts. In conclusion, figure-ground activity in V1 may act as an intermediate component linking visual and motor signals.

Animals↗

Learning receptive fields using predictive feedback.

Previously, it was suggested that feedback connections from higher- to lower-level areas carry predictions of lower-level neural activities, whereas feedforward connections carry the residual error between the predictions and the actual lower-level activities [Rao, R.P.N., Ballard, D.H., 1999. Nature Neuroscience 2, 79-87.]. A computational model implementing the hypothesis learned simple cell receptive fields when exposed to natural images. Here, we use predictive feedback to explain tuning properties in medial superior temporal area (MST). We implement the hypothesis using a new, biologically plausible, algorithm based on matching pursuit, which retains all the features of the previous implementation, including its ability to efficiently encode input. When presented with natural images, the model developed receptive field properties as found in primary visual cortex. In addition, when exposed to visual motion input resulting from movements through space, the model learned receptive field properties resembling those in MST. These results corroborate the idea that predictive feedback is a general principle used by the visual system to efficiently encode natural input.

Algorithms↗

Anatomical and functional MR imaging in the macaque monkey using a vertical large-bore 7 Tesla setup.

Functional magnetic resonance imaging (MRI) in the nonhuman primate promises to provide a much desired link between brain research in humans and the large body of systems neuroscience work in animals. We present here a novel high field, large-bore, vertical MR system (7 T/60 cm, 300 MHz), which was optimized for neuroscientific research in macaque monkeys. A strong magnetic field was applied to increase sensitivity and spatial resolution for both MRI and spectroscopy. Anatomical imaging with voxel sizes as small as 75x150x300 microm3 and with high contrast-to-noise ratios permitted the visualization of the characteristic lamination of some neocortical areas, e.g., Baillarger lines. Relaxation times were determined for different structures: at 7 T, T1 was 2.01/1.84/1.54 s in GM/GM-V1/WM, T2 was 59.1/54.4 ms in GM/WM and T2* was 29 ms. At 4.7 T, T1 was 25% shorter, T2 and T2* 18% longer compared to 7T. Spatiotemporally resolved blood-oxygen-level-dependent (BOLD) signal changes yielded robust activations and deactivations (negative BOLD), with average amplitudes of 4.1% and -2.4%, respectively. Finally, the first high-resolution (500 microm in-plane) images of cerebral blood flow in the anesthetized monkey are presented. On functional activation we observed flow increases of up to 38% (59 to 81 ml/100 g/min) in the primary visual cortex, V1. Compared to BOLD maps, functional CBF maps were found to be localized entirely within the gray matter, providing unequivocal evidence for high spatial specificity. The exquisite sensitivity of the system and the increased specificity of the hemodynamic signals promise further insights into the relationship of the latter to the underlying physiological activity.

Animals↗

Development of visually evoked cortical activity in infant macaque monkeys studied longitudinally with fMRI.

We studied the development of visual activation longitudinally in two infant monkeys aged 103-561 days using the BOLD fMRI technique under opiate anesthesia and compared the results with those obtained in three adult animals studied under identical conditions. Visual activation in primary visual cortex, V1, was strong and reliable in monkeys of the youngest and oldest ages, showing that functional imaging techniques give qualitatively similar results in infants and adults. Visual activation in extrastriate areas involved in processing motion (MT/V5) and form (V4) was not evident in the younger animals, but became more adult-like in the older animals. This delayed onset of measurable BOLD responses in extrastriate visual cortex may reflect delayed development of visual responses in these areas, although at this stage it is not possible to rule out either effects of anesthesia or of changes in cerebral vascular response mechanisms as the cause. The demonstration of visually evoked BOLD responses in young monkeys shows that the BOLD fMRI technique can usefully be employed to address functional questions of brain development.

Animals↗

Laminar specificity in monkey V1 using high-resolution SE-fMRI.

The lamination of mammalian neocortex is widely used as reference for describing a wide range of anatomical and physiological data. Its value lies in the observation that in all examined species, cortical afferents, intrinsic cells and projection neurons organize themselves with respect to the laminae. The comprehension of the computations, carried out by the neocortical microcircuits, critically relies on the study of the interlaminar connectivity patterns and the intralaminar physiological processes in vivo. High-resolution functional neuroimaging, enabling the visualization of activity in individual cortical laminae or columns, may greatly contribute in such studies. Yet, the BOLD effect, as measured with the commonly used GE-EPI, contains contributions from both macroscopic venous blood vessels and capillaries. The low density of the cortical veins limits the effective spatial specificity of the fMRI signal and yields maps that are weighted toward the macrovasculature, which thus can be significantly different from the actual site of increased neuronal activity. Spin-echo (SE) sequences yielding apparent T2-weighted BOLD images have been shown to improve spatial specificity by increasing the sensitivity of the signal to spins of the parenchyma, particularly at high magnetic fields. Here we used SE-fMRI at 4.7 T to examine the specificity and resolution of functional maps obtained by stimulating the primary visual cortex of monkeys. Cortical layers could be clearly visualized, and functional activity was predominantly localized in cortical layer IV/Duvernoy layer 3. The choice of sequence parameters influences the fMRI signal, as the SE-EPI is by nature sensitive to T2* in addition to its T2 dependency. Using parameters that limit T2* effects yielded higher specificity and better visualization of the cortical laminae. Because the demands of high-spatial resolution using SE severely decreases temporal resolution, we used a stimulus protocol that allows sampling at higher effective temporal resolution. This way, it was possible to acquire high-spatial and high-temporal resolution SE-fMRI data.

Animals↗

A role for the 'magnocellular advantage' in visual impairments in neurodevelopmental and psychiatric disorders.

Evidence exists implicating abnormal visual information processing and visually driven attention in a number of neurodevelopmental and psychiatric disorders, suggesting that research into such disorders may benefit from a better understanding of more recent advances in visual system processing. A new integrated model of visual processing based on primate single cell and human electrophysiology may provide a framework, to understand how the visual system is involved, by implicating the magnocellular pathway's role in driving attentional mechanisms in higher-order cortical regions, what we term the 'magnocellular advantage'. Evidence is also presented demonstrating visual processing occurs considerably faster than previously assumed, and emphasising the importance of top-down feedback signals into primary visual cortex, as well as considering the possibility of lateral connections from dorsal to ventral visual areas. Such organisation is argued to be important for future research highlighting visual aspects of impairment in disorders as diverse as schizophrenia and autism.

Animals↗

Naso-temporal asymmetry of the N170 for processing faces in normal viewers but not in developmental prosopagnosia.

Some elementary aspects of faces can be processed before cortical maturation or after lesion of primary visual cortex. Recent findings suggesting a role of an evolutionary ancient visual system in face processing have exploited the relative advantage of the temporal hemifield (nasal hemiretina). Here, we investigated whether under some circumstances face processing also shows a temporal hemifield advantage. We measured the face sensitive N170 to laterally presented faces viewed passively under monocular conditions and compared face recognition in the temporal and nasal hemiretina. A N170 response for upright faces was observed which was larger for projections to the nasal hemiretina/temporal hemifields. This pattern was not observed in a developmental prosopagnosic. These results point to the importance of the early stages of face processing for normal face recognition abilities and suggest a potentially important factor in the origins of developmental prosopagnosia.

Adult↗

SMI-32 parcellates the visual cortical areas of the marmoset.

The distribution pattern of SMI-32-immunoreactivity (SMI-32-ir) of neuronal elements was examined in the visual cortical areas of marmoset monkey. Layer IV of the primary visual cortex (V1) and layers III and V of the extrastriate areas showed the most abundant SMI-32-ir. The different areal and laminar distribution of SMI-32-ir allowed the distinction between various extrastriate areas and determined their exact anatomical boundaries in the New World monkey, Callithrix penicillata. It is shown here that the parcellating nature of SMI-32 described earlier in the visual cortical areas of other mammals - including Old World monkeys - is also present in the marmoset. Furthermore, a comparison became possible between the chemoanatomical organization of New World and Old World primates' visual cortical areas.

Animals↗

Visual attention deficits in Alzheimer's disease: an fMRI study.

Cognitive and neuroscience studies indicate that attentional operations are impaired in Alzheimer's disease (AD). Our goal was to define the anatomical areas of activation associated with visual attention processing and to define deficits or changes that may occur in AD patients compared with control group. Thirteen AD patients and 13 age- and education-matched normal controls were tested in two visual search tasks (one was a conjunction task, where feature binding is required. The other was a subset task, where group stimuli is needed without feature binding) using fMRI techniques. After stereotactical normalization, voxel-by-voxel t statistics was used to compare activated brain areas between patients and control subjects. Our findings suggest that both search tasks are controlled by partially overlapping cerebral networks, including parietal, frontal and occipital-temporal cortical regions and primary visual cortex. The AD patient group showed less activation in both parietal lobes and the left frontal regions, while increased activation was found in the right frontal lobes and the right occipito-temporal cortical regions with the conjunction task. In the subset task, decreased activation in AD patients was seen in the left parietal lobe and bilateral frontal lobes, while increased activation was seen in both medial temporal lobes. In addition, for the comparison between tasks, The difference is very small for AD patients. Control group showed a higher amplitude in the right prefrontal region, temporal cortical regions and parietal lobe. These results indicate that attention deficits in AD patients may be attributed to both binding problem and grouping inefficiency.

Aged↗

Volatile anesthetics disrupt frontal-posterior recurrent information transfer at gamma frequencies in rat.

We seek to understand neural correlates of anesthetic-induced unconsciousness. We hypothesize that cortical integration of sensory information may underlie conscious perception and may be disrupted by anesthetics. A critical role in frontal-posterior interactions has been proposed, and gamma (20-60 Hz) oscillations have also been assigned an essential role in consciousness. Here we investigated whether general anesthetics may interfere with the exchange of information encoded in gamma oscillations between frontal and posterior cortices. Bipolar electrodes for recording of event-related potentials (ERP) were chronically implanted in the primary visual cortex, parietal association and frontal association cortices of six rats. Sixty light flashes were presented every 5s, and ERPs were recorded at increasing concentrations of halothane or isoflurane (0-2%). Information exchange was estimated by transfer entropy, a novel measure of directional information transfer. Transfer entropy was calculated from 1-s wavelet-transformed ERPs. We found that (1) feedforward transfer entropy (FF-TE) and feedback transfer entropy (FB-TE) were balanced in conscious-sedated state; (2) anesthetics at concentrations producing unconsciousness augmented both FF-TE and FB-TE at 30 Hz but reduced them at 50 Hz; (3) reduction at 50 Hz was more pronounced for FB-TE, especially between frontal and posterior regions; (4) at high concentrations, both FF-TE and FB-TE at all frequencies were at or below conscious-sedated baseline. Our findings suggest that inhalational anesthetics preferentially impair frontal-posterior FB information transfer at high gamma frequencies consistent with the postulated role of frontal-posterior interactions in consciousness.

Anesthetics, Inhalation↗

Diffusion tensor imaging in acquired blind humans.

Retinal implants as a future possible therapy of blindness rely on an intact neural transmission from the retina to the primary visual cortex. By now it remains unknown, in how far the absence of afferent input in blindness affects also the organization of the optic radiation. Using diffusion tensor imaging (DTI), the non-invasive evaluation of large fiber tracts including the optic radiation has become possible. This method is sensitive to changes of the axonal state such as wallerian degeneration. We have compared DTI data from 6 acquired blind patients with those of a group of 11 healthy control subjects. Neither the relative anisotropy quotient of the visual fiber tract and the pyramidal tract showed a statistically significant difference between the blind patients and the control group nor did the absolute values of the relative anisotropy in the pyramidal tract and the visual fiber tract. There was no axonal degeneration of the optic radiation in late onset acquired blindness. With the optic pathways remaining intact, transmitting electric signals of retinal implants to the visual regions of the human brain seems to be possible even after decades of acquired blindness.

Adult↗

Associative learning in early vision.

Sensory discriminations often improve with practice (perceptual learning). Recent results show that practice does not necessarily lead to the best possible performance on the task. It was shown that learning a task (contrast discrimination) that has already reached saturation could be enabled by a contextual change in the stimulus (the addition of surrounding flankers) during practice. Psychophysical results with varying context show a behavior that is described by a network of local visual processors with horizontal recurrent interactions. We describe a mathematical learning rule for the modification of cortical synapses that is inspired by the experimental results and apply it to recurrent cortical networks that respond to external stimuli. The model predicts that repeated presentation of the same stimulus leads to saturation of synaptic modification, such that the strengths of recurrent connections depend on the configuration of the stimulus but not on its amplitude. When a new stimulus is introduced, the modification is rekindled until a new equilibrium is reached. This effect may explain the saturation of perceptual learning when practicing a certain task repeatedly. We present simulations of contrast discrimination in a simplified model of a cortical column in the primary visual cortex and show that performance of the model is reminiscent of context-dependent perceptual learning.

Computer Simulation↗

Nonlinear V1 responses to natural scenes revealed by neural network analysis.

A key goal in the study of visual processing is to obtain a comprehensive description of the relationship between visual stimuli and neuronal responses. One way to guide the search for models is to use a general nonparametric regression algorithm, such as a neural network. We have developed a multilayer feed-forward network algorithm that can be used to characterize nonlinear stimulus-response mapping functions of neurons in primary visual cortex (area V1) using natural image stimuli. The network is capable of extracting several known V1 response properties such as: orientation and spatial frequency tuning, the spatial phase invariance of complex cells, and direction selectivity. We present details of a method for training networks and visualizing their properties. We also compare how well conventional explicit models and those developed using neural networks can predict novel responses to natural scenes.

Action Potentials↗

Perceptual grouping and the interactions between visual cortical areas.

Visual perception involves the grouping of individual elements into coherent patterns, such as object representations, that reduce the descriptive complexity of a visual scene. The computational and physiological bases of this perceptual remain poorly understood. We discuss recent fMRI evidence from our laboratory where we measured activity in a higher object processing area (LOC), and in primary visual cortex (V1) in response to visual elements that were either grouped into objects or randomly arranged. We observed significant activity increases in the LOC and concurrent reductions of activity in V1 when elements formed coherent shapes, suggesting that activity in early visual areas is reduced as a result of grouping processes performed in higher areas. In light of these results we review related empirical findings of context-dependent changes in activity, recent neurophysiology research related to cortical feedback, and computational models that incorporate feedback operations. We suggest that feedback from high-level visual areas reduces activity in lower areas in order to simplify the description of a visual image-consistent with both predictive coding models of perception and probabilistic notions of 'explaining away.'

Animals↗

Homeostatic synaptic scaling in self-organizing maps.

Various forms of the self-organizing map (SOM) have been proposed as models of cortical development [Choe Y., Miikkulainen R., (2004). Contour integration and segmentation with self-organized lateral connections. Biological Cybernetics, 90, 75-88; Kohonen T., (2001). Self-organizing maps (3rd ed.). Springer; Sirosh J., Miikkulainen R., (1997). Topographic receptive fields and patterned lateral interaction in a self-organizing model of the primary visual cortex. Neural Computation, 9(3), 577-594]. Typically, these models use weight normalization to contain the weight growth associated with Hebbian learning. A more plausible mechanism for controlling the Hebbian process has recently emerged. Turrigiano and Nelson [Turrigiano G.G., Nelson S.B., (2004). Homeostatic plasticity in the developing nervous system. Nature Reviews Neuroscience, 5, 97-107] have shown that neurons in the cortex actively maintain an average firing rate by scaling their incoming weights. In this work, it is shown that this type of homeostatic synaptic scaling can replace the common, but unsupported, standard weight normalization. Organized maps still form and the output neurons are able to maintain an unsaturated firing rate, even in the face of large-scale cell proliferation or die-off. In addition, it is shown that in some cases synaptic scaling leads to networks that more accurately reflect the probability distribution of the input data.

Animals↗

Intrinsic signal recording from a monkey whose behavior was maintained by a schedule of reinforcement.

Optical recording of cortical activity in awake monkeys has enhanced our understanding of the functional anatomy of the primary visual cortex (V1). However, cortical representation of visual cognition has not been studied by optical recording, even though the greatest merit of using awake animals is that they can offer advantages in studying cognitive function that anesthetized animals cannot. Thus far, the optical recording method has not been combined with tasks that accompany body movements because of concern about movement noise, although behavioral tasks are helpful in the study of animal cognition. Here, I tested the influence of body movements during the signal acquisition period on the resultant images. I recorded the intrinsic signals associating with different orientations from V1 of a monkey who was emitting behavior during the signal acquisition period. Although the monkey's behavior was maintained on a variable-interval schedule that typically induces a high rate of response, orientation maps were consistently obtained. Therefore, a successful recording under this operant regimen implies the applicability of the optical recording method to other behavioral tasks. Several constraints in applying optical recording to studies using behaving animals are also discussed.

Analysis of Variance↗