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Christof Koch

Publications and source records attributed to Christof Koch.

At least 37 records · Page 2Linked to original sources

Components of bottom-up gaze allocation in natural images.

Recent research [Parkhurst, D., Law, K., & Niebur, E., 2002. Modeling the role of salience in the allocation of overt visual attention. Vision Research 42 (1) (2002) 107-123] showed that a model of bottom-up visual attention can account in part for the spatial locations fixated by humans while free-viewing complex natural and artificial scenes. That study used a definition of salience based on local detectors with coarse global surround inhibition. Here, we use a similar framework to investigate the roles of several types of non-linear interactions known to exist in visual cortex, and of eccentricity-dependent processing. For each of these, we added a component to the salience model, including richer interactions among orientation-tuned units, both at spatial short range (for clutter reduction) and long range (for contour facilitation), and a detailed model of eccentricity-dependent changes in visual processing. Subjects free-viewed naturalistic and artificial images while their eye movements were recorded, and the resulting fixation locations were compared with the models' predicted salience maps. We found that the proposed interactions indeed play a significant role in the spatiotemporal deployment of attention in natural scenes; about half of the observed inter-subject variance can be explained by these different models. This suggests that attentional guidance does not depend solely on local visual features, but must also include the effects of interactions among features. As models of these interactions become more accurate in predicting behaviorally-relevant salient locations, they become useful to a range of applications in computer vision and human-machine interface design.

Adolescent↗

Intrinsic noise in cultured hippocampal neurons: experiment and modeling.

Ion channels open and close stochastically. The fluctuation of these channels represents an intrinsic source of noise that affects the input-output properties of the neuron. We combined whole-cell measurements with biophysical modeling to characterize the intrinsic stochastic and electrical properties of single neurons as observed at the soma. We measured current and voltage noise in 18 d postembryonic cultured neurons from the rat hippocampus, at various subthreshold and near-threshold holding potentials in the presence of synaptic blockers. The observed current noise increased with depolarization, as ion channels were activated, and its spectrum demonstrated generalized 1/f behavior. Exposure to TTX removed a significant contribution from Na+ channels to the noise spectrum, particularly at depolarized potentials, and the resulting spectrum was now dominated by a single Lorentzian (1/f2) component. By replacing the intracellular K+ with Cs+, we demonstrated that a major portion of the observed noise was attributable to K+ channels. We compared the measured power spectral densities to a 1-D cable model of channel fluctuations based on Markov kinetics. We found that a somatic compartment, in combination with a single equivalent cylinder, described the effective geometry from the viewpoint of the soma. Four distinct channel populations were distributed in the membrane and modeled as Lorentzian current noise sources. Using the NEURON simulation program, we summed up the contributions from the spatially distributed current noise sources and calculated the total voltage and current noise. Our quantitative model reproduces important voltage- and frequency-dependent features of the data, accounting for the 1/f behavior, as well as the effects of various blockers.

Animals↗

Qualia.

Explore the source record for details and available documents.

Brain↗

Face-gender discrimination is possible in the near-absence of attention.

The attentional cost associated with the visual discrimination of the gender of a face was investigated. Participants performed a face-gender discrimination task either alone (single-task) or concurrently (dual-task) with a known attentional demanding task (5-letter T/L discrimination). Overall performance on face-gender discrimination suffered remarkably little under the dual-task condition compared to the single-task condition. Similar results were obtained in experiments that controlled for potential training effects or the use of low-level cues in this discrimination task. Our results provide further evidence against the notion that only low-level representations can be accessed outside the focus of attention.

Adult↗

Motion-induced blindness does not affect the formation of negative afterimages.

Aftereffects induced by invisible stimuli constitute a powerful tool to investigate what type of neural information processing can occur in the absence of visual awareness. This approach has been successfully used to demonstrate that awareness of oriented gratings or translating stimuli is not necessary to obtain a robust orientation-specific or motion-specific aftereffect. We exploit motion-induced blindness (MIB, Bonneh, Cooperman, & Sagi, 2001) to investigate the related question of the influence of visual awareness on the formation of negative afterimages. Our results show that MIB does not affect the persistence and intensity of afterimages. Thus, there is no significant contribution to the formation of afterimages beyond the sites mediating MIB.

Adult↗

A learning rule for local synaptic interactions between excitation and shunting inhibition.

The basic requirement for direction selectivity is a nonlinear interaction between two different inputs in space-time. In some models, the interaction is hypothesized to occur between excitation and inhibition of the shunting type in the neuron's dendritic tree. How can the required spatial specificity be acquired in an unsupervised manner? We here propose an activity-based, local learning model that can account for direction selectivity in visual cortex based on such a local veto operation and that depends on synaptically induced changes in intracellular calcium concentration. Our biophysical simulations suggest that a model cell with our learning algorithm can develop direction selectivity organically after unsupervised training. The learning rule is also applicable to a neuron with multiple-direction-selective subunits and to a pair of cells with opposite-direction selectivities and is stable under different starting conditions, delays, and velocities.

Algorithms↗

Consciousness and neurosurgery.

The neuronal basis of consciousness is the greatest challenge to the scientific worldview. Much relevant empirical work is carried out on the minimal neuronal mechanisms underlying any one specific conscious percept. Two broad approaches are popular among brain scientists: electrophysiological recordings from individual neurons in the cortex of behaving monkeys or behavior combined with functional brain imaging in humans. However, many aspects of consciousness are problematic or remain off-limits to the former approach, while the latter one lacks sufficient spatial and temporal resolution to monitor individual neurons that are key to perception, thought, memory, and action. It is here that neurosurgeons, probing the living human brain on a daily basis, can play a decisive role. This article explores the contributions of neurosurgeons to this quest and outlines some of the results that have already been achieved.

Animals↗

Trace but not delay fear conditioning requires attention and the anterior cingulate cortex.

Higher cognitive functions such as attention have been difficult to model in genetically tractable organisms. In humans, attention-distracting stimuli interfere with trace but not delay conditioning, two forms of associative learning. Attention has also been correlated with activation of anterior cingulate cortex (ACC), but its functional significance is unclear. Here we show that a visual distractor interferes selectively with trace but not delay auditory fear conditioning in mice. Trace conditioning is associated with increased neuronal activity in ACC, as assayed by relative levels of c-fos expression, and is selectively impaired by lesions of this structure. The effects of the ACC lesions are unlikely to be caused by indirect impairment of the hippocampus, which is required for mnemonic aspects of trace conditioning. These data suggest that trace conditioning may be useful for studying neural substrates of attention in mice, and implicate the ACC as one such substrate.

Animals↗

Visual selective behavior can be triggered by a feed-forward process.

The ventral visual pathway implements object recognition and categorization in a hierarchy of processing areas with neuronal selectivities of increasing complexity. The presence of massive feedback connections within this hierarchy raises the possibility that normal visual processing relies on the use of computational loops. It is not known, however, whether object recognition can be performed at all without such loops (i.e., in a purely feed-forward mode). By analyzing the time course of reaction times in a masked natural scene categorization paradigm, we show that the human visual system can generate selective motor responses based on a single feed-forward pass. We confirm these results using a more constrained letter discrimination task, in which the rapid succession of a target and mask is actually perceived as a distractor. We show that a masked stimulus presented for only 26 msec - and often not consciously perceived - can fully determine the earliest selective motor responses: The neural representations of the stimulus and mask are thus kept separated during a short period corresponding to the feed-forward "sweep." Therefore, feedback loops do not appear to be "mandatory" for visual processing. Rather, we found that such loops allow the masked stimulus to reverberate in the visual system and affect behavior for nearly 150 msec after the feed-forward sweep.

Attention↗

Working memory and fear conditioning.

Previous studies of associative learning implicate higher-level cognitive processes in some forms of classical conditioning. An ongoing debate is concerned with the extent to which attention and awareness are necessary for trace but not delay eye-blink conditioning [Clark, R. E. & Squire, L. R. (1998) Science 280, 77-81; Lovibond, P. F. & Shanks, D. (2002) J. Exp. Psychol. Anim. Behav. Processes 28, 38-42]. In trace conditioning, a short interval is interposed between the termination of the conditioned stimulus (CS) and the onset of the unconditioned stimulus (US). In delay conditioning, the CS and US overlap. We here investigate the extent to which human classical fear conditioning depends on working memory. Subjects had to carry out an n-back task, requiring tracking an item 1 or 2 back in a sequentially presented list of numbers, while simultaneously being tested for their ability to associate auditory cues with shocks under a variety of conditions (single-cue versus differential; delay versus trace; no task versus 0-, 1-, and 2-back). Differential delay conditioning proved to be more resilient than differential trace conditioning but does show a reduction due to task interference similar in slope to that found in trace conditioning. Explicit knowledge of the stimulus contingency facilitates but does not guarantee trace conditioning. Only the single-cue delay protocol shows conditioning during the more difficult working memory task. Our findings suggest that the larger the cognitive demands on the system, the less likely conditioning occurs. A postexperimental questionnaire showed a positive correlation between conditioning and awareness for differential trace conditioning extinction.

Adolescent↗

Fusion of competing features is not serial.

How features of an object are bound into a unique percept is one of the puzzling problems in the cognitive and neuro-sciences. In order to investigate the spatio-temporal mechanisms of feature binding, we serially present two verniers with opposite offset directions for very short durations. Only one vernier is perceived with its offset dominated by the vernier presented second. This dominance reverses if the two verniers are followed by masking gratings, i.e. the first presented vernier dominates performance. Therefore, feature fusion can neither be explained completely by spatially local mechanisms nor by the temporal order of appearance of elements.

Adult↗

Human visual object categorization can be described by models with low memory capacity.

Studies of high-level models of visual object categorization have left unresolved issues of neurobiological relevance, including how features are extracted from the image and the role played by memory capacity in categorization performance. We compared the ability of a comprehensive set of models to match the categorization performance of human observers while explicitly accounting for the models' numbers of free parameters. The most successful models did not require a large memory capacity, suggesting that a sparse, abstracted representation of category properties may underlie categorization performance. This type of representation--different from classical prototype abstraction--could also be extracted directly from two-dimensional images via a biologically plausible early-vision model, rather than relying on experimenter-imposed features.

Adolescent↗

Is perception discrete or continuous?

How does conscious perception evolve following stimulus presentation? The idea that perception relies on discrete processing epochs has been often considered, but never widely accepted. The alternative, a continuous translation of the external world into explicit perception, although more intuitive and subjectively appealing, cannot satisfactorily account for a large body of psychophysical data. Cortical and thalamocortical oscillations in different frequency bands could provide a neuronal basis for such discrete processes, but are rarely analyzed in this context. This article reconciles the unduly abandoned topic of discrete perception with current views and advances in neuroscience.

Journal Article↗

A framework for consciousness.

Here we summarize our present approach to the problem of consciousness. After an introduction outlining our general strategy, we describe what is meant by the term 'framework' and set it out under ten headings. This framework offers a coherent scheme for explaining the neural correlates of (visual) consciousness in terms of competing cellular assemblies. Most of the ideas we favor have been suggested before, but their combination is original. We also outline some general experimental approaches to the problem and, finally, acknowledge some relevant aspects of the brain that have been left out of the proposed framework.

Action Potentials↗

Modeling reverse-phi motion-selective neurons in cortex: double synaptic-veto mechanism.

Reverse-phi motion is the illusory reversal of perceived direction of movement when the stimulus contrast is reversed in successive frames. Livingstone, Tsao, and Conway (2000) showed that direction-selective cells in striate cortex of the alert macaque monkey showed reversed excitatory and inhibitory regions when two different contrast bars were flashed sequentially during a two-bar interaction analysis. While correlation or motion energy models predict the reverse-phi response, it is unclear how neurons can accomplish this. We carried out detailed biophysical simulations of a direction-selective cell model implementing a synaptic shunting scheme. Our results suggest that a simple synaptic-veto mechanism with strong direction selectivity for normal motion cannot account for the observed reverse-phi motion effect. Given the nature of reverse-phi motion, a direct interaction between the ON and OFF pathway, missing in the original shunting-inhibition model, it is essential to account for the reversal of response. We here propose a double synaptic-veto mechanism in which ON excitatory synapses are gated by both delayed ON inhibition at their null side and delayed OFF inhibition at their preferred side. The converse applies to OFF excitatory synapses. Mapping this scheme onto the dendrites of a direction-selective neuron permits the model to respond best to normal motion in its preferred direction and to reverse-phi motion in its null direction. Two-bar interaction maps showed reversed excitation and inhibition regions when two different contrast bars are presented.

Animals↗

Competition and selection during visual processing of natural scenes and objects.

When a visual scene, containing many discrete objects, is presented to our retinae, only a subset of these objects will be explicitly represented in visual awareness. The number of objects accessing short-term visual memory might be even smaller. Finally, it is not known to what extent "ignored" objects (those that do not enter visual awareness) will be processed--or recognized. By combining free recall, forced-choice recognition and visual priming paradigms for the same natural visual scenes and subjects, we were able to estimate these numbers, and provide insights as to the fate of objects that are not explicitly recognized in a single fixation. When presented for 250 ms with a scene containing 10 distinct objects, human observers can remember up to 4 objects with full confidence, and between 2 and 3 more when forced to guess. Importantly, the objects that the subjects consistently failed to report elicited a significant negative priming effect when presented in a subsequent task, suggesting that their identity was represented in high-level cortical areas of the visual system, before the corresponding neural activity was suppressed during attentional selection. These results shed light on neural mechanisms of attentional competition, and representational capacity at different levels of the human visual system.

Attention↗

Multiplicative computation in a visual neuron sensitive to looming.

Multiplicative operations are important in sensory processing, but their biophysical implementation remains largely unknown. We investigated an identified neuron (the lobula giant movement detector, LGMD, of locusts) whose output firing rate in response to looming visual stimuli has been described by two models, one of which involves a multiplication. In this model, the LGMD multiplies postsynaptically two inputs (one excitatory, one inhibitory) that converge onto its dendritic tree; in the other model, inhibition is presynaptic to the LGMD. By using selective activation and inactivation of pre- and postsynaptic inhibition, we show that postsynaptic inhibition has a predominant role, suggesting that multiplication is implemented within the neuron itself. Our pharmacological experiments and measurements of firing rate versus membrane potential also reveal that sodium channels act both to advance the response of the LGMD in time and to map membrane potential to firing rate in a nearly exponential manner. These results are consistent with an implementation of multiplication based on dendritic subtraction of two converging inputs encoded logarithmically, followed by exponentiation through active membrane conductances.

Action Potentials↗

Rapid natural scene categorization in the near absence of attention.

What can we see when we do not pay attention? It is well known that we can be "blind" even to major aspects of natural scenes when we attend elsewhere. The only tasks that do not need attention appear to be carried out in the early stages of the visual system. Contrary to this common belief, we report that subjects can rapidly detect animals or vehicles in briefly presented novel natural scenes while simultaneously performing another attentionally demanding task. By comparison, they are unable to discriminate large T's from L's, or bisected two-color disks from their mirror images under the same conditions. We conclude that some visual tasks associated with "high-level" cortical areas may proceed in the near absence of attention.

Adult↗