Biomedical subjects

Norberto M Grzywacz

Publications and source records attributed to Norberto M Grzywacz.

16 recordsLinked to original sources

PubMed2006

Estimating nonlinear receptive fields from natural images.

The response of visual cells is a nonlinear function of their stimuli. In addition, an increasing amount of evidence shows that visual cells are optimized to process natural images. Hence, finding good nonlinear models to characterize visual cells using natural stimuli is important. The Volterra model is an appealing nonlinear model for visual cells. However, their large number of parameters and the limited size of physiological recordings have hindered its application. Recently, a substantiated hypothesis stating that the responses of each visual cell could depend on an especially low-dimensional subspace of the image space has been proposed. We use this low-dimensional subspace in the Volterra relevant-space technique to allow the estimation of high-order Volterra models. Most laboratories characterize the response of visual cells as a nonlinear function on the low-dimensional subspace. They estimate this nonlinear function using histograms and by fitting parametric functions to them. Here, we compare the Volterra model with these histogram-based techniques. We use simulated data from cortical simple cells as well as simulated and physiological data from cortical complex cells. Volterra models yield equal or superior predictive power in all conditions studied. Several methods have been proposed to estimate the low-dimensional subspace. In this article, we test projection pursuit regression (PPR), a nonlinear regression algorithm. We compare PPR with two popular models used in vision: spike-triggered average (STA) and spike-triggered covariance (STC). We observe that PPR has advantages over these alternative algorithms. Hence, we conclude that PPR is a viable algorithm to recover the relevant subspace from natural images and that the Volterra model, estimated through the Volterra relevant-space technique, is a compelling alternative to histogram-based techniques.

Algorithms↗

PubMed2006

Modeling Starburst cells' GABA(B) receptors and their putative role in motion sensitivity.

Neal and Cunningham (Neal, M. J., and J. R. Cunningham. 1995. J. Physiol. (Lond.). 482:363-372) showed that GABA(B) agonists and glycinergic antagonists enhance the light-evoked release of retinal acetylcholine. They proposed that glycinergic cells inhibit the cholinergic Starburst amacrine cells and are in turn inhibited by GABA through GABA(B) receptors. However, as recently shown, glycinergic cells do not appear to have GABA(B) receptors. In contrast, the Starburst amacrine cell has GABA(B) receptors in a subpopulation of its varicosities. We thus propose an alternate model in which GABA(B)-receptor activation reduces the release of ACh from some dendritic compartments onto a glycinergic cell, which then feeds back and inhibits the Starburst cell. In this model, the GABA necessary to make these receptors active comes from the Starburst cell itself, making them autoreceptors. Computer simulations of this model show that it accounts quantitatively for the Neal and Cunningham data. We also argue that GABA(B) receptors could work to increase the sensitivity to motion over other stimuli.

Acetylcholine↗

PubMed2006

AII amacrine cells in the distal inner nuclear layer of the mouse retina.

We serendipitously found a distal Disabled-1 (Dab1)-immunoreactive cell in retina of the C57BL/6J black mouse. The somata of these cells are located in the outermost part of the inner nuclear layer (INL). Their processes extend toward the outer plexiform layer (OPL), receiving synaptic inputs from horizontal and interplexiform cells. In the current study, we name this cell the "distal Dab1-immunoreactive cell." Double-labeling experiments demonstrate that the distal Dab1-immunoreactive cell is not a horizontal cell. Rather, the distal Dab1 cell appears to be a misplaced AII cell, by being glycine transporter-1-immunoreactive and by resembling the latter cell in an electron microscopic analysis. A distal Dab1 cell had been reported in the FVB/N mouse retina, a model of retinitis pigmentosa (Park et al. [2004] Cell Tissue Res 315:407-412). However, here, we found this distal Dab1-immunoreactive cell in the adult and normal developing mouse retinas. Hence, we show that such cells do not require the loss of photoreceptors as suggested previously (Park et al. [2004] Cell Tissue Res 315:407-412). Instead, two other pieces of data suggest an alternative explanation sources for distal Dab1 cells. First, we find a correlation between the number of these cells in the left and right eyes Second, developmental analysis shows that the distal Dab1-immunoreactive cell is first observed shortly after birth. At the same time, AII cells emerge, extending their neurites into the inner retina. These data suggest that distal Dab1-immunoreactive cells are misplaced AII amacrine cells, resulting from genetically modulated anomalies owing to migration errors.

Amacrine Cells↗

PubMed2006

Statistically robust detection of spontaneous, non-stereotypical neural signals.

Neural signals of interest are often temporally spontaneous and non-stereotypical in waveform. Detecting such signals is difficult, since one cannot use time-locking or simple template-matching techniques. We have sought a statistical method for automatically estimating the baseline in these conditions, and subsequently detecting the occurrence of neural signals. One could consider the signals as outliers in the distribution of neural activity and thus separate them from the baseline with median-based techniques. However, we found that baseline estimators that rely on the median are problematic. They introduce progressively greater estimation errors as the neural signal's duration, amplitude or frequency increases. Therefore, we tested several mode-based algorithms, taking advantage of the most probable state of the neural activity being the baseline. We found that certain mode-based algorithms perform baseline estimation well, with low susceptibility to changes in event duration, amplitude or frequency. Once the baseline is properly established, its median absolute deviation (MAD) can be determined. One can then use it to detect spontaneous signals robustly as outliers from the noise distribution. We also demonstrate how the choice of detection threshold in terms of MADs can be used to bias against false positives, without creating too many false negatives or vice versa.

Action Potentials↗

PubMed2006

Dark-rearing-induced reduction of GABA and GAD and prevention of the effect by BDNF in the mouse retina.

Gamma-aminobutyric acid (GABA) is an important retinal neurotransmitter. We studied the expression of GABA, glutamate decarboxylase 65 (GAD65) and GAD67 by immunocytochemistry and Western blot, in the retinas of control and dark-reared C57BL/6J black mice. This study asked three questions. First, is visual input necessary for the normal expression of GABA, GAD65 and GAD67? Second, can the retina recover from the effects of dark-rearing if returned to a normal light-dark cycle? Third, does BDNF prevent the influence of dark-rearing on the expression of GABA and GAD? At postnatal day 10 (P10), before eye opening, GABA immunoreactivity was present in the ganglion cell layer (GCL), in the innermost rows of the inner nuclear layer (INL) and throughout the inner plexiform layer (IPL) of control and dark-reared retinas. In P30 control retinas, GABA immunoreactivity showed similar patterns to those at P10. However, in P30 dark-reared retinas, the density of GABA-immunoreactive cells was lower in both the INL and GCL than in control retinas. In addition, visual deprivation retarded GABA immunoreactivity in the IPL. Western blot analysis showed corresponding differences in the levels of GAD65 but not of GAD67 expression between control and dark-rearing conditions. In our study, dark-rearing effects were reversed when the mice were put in normal cyclic light-dark conditions for 2 weeks. Moreover, dark-reared retinas treated with BDNF showed normal expression of both GABA and GAD65. Our data indicate that normal expression of GABA and GAD65 is dependent on visual input. Furthermore, the data suggest that BDNF controls this dependence.

Animals↗

PubMed2005

Compartmental localization of gamma-aminobutyric acid type B receptors in the cholinergic circuitry of the rabbit retina.

Although many effects of gamma-aminobutyric acid (GABA) on retinal function have been attributed to GABA(A) and GABA(C) receptors, specific retinal functions have also been shown to be mediated by GABA(B) receptors, including facilitation of light-evoked acetylcholine release from the rabbit retina (Neal and Cunningham [1995] J. Physiol. 482:363-372). To explain the results of a rich set of experiments, Neal and Cunningham proposed a model for this facilitation. In this model, GABA(B) receptor-mediated inhibition of glycinergic cells would reduce their own inhibition of cholinergic cells. In turn, muscarinic input from the latter to the glycinergic cells would complete a negative-feedback circuitry. In this study, we have used immunohistochemical techniques to test elements of this model. We report that glycinergic amacrine cells are GABA(B) receptor negative. In contrast, our data reveal the localization of GABA(B) receptors on cholinergic/GABAergic starburst amacrine cells. High-resolution localization of GABA(B) receptors on starburst amacrine cells shows that they are discretely localized to a limited population of its varicosities, the majority of likely synaptic-release sites being devoid of detectable levels of GABA(B) receptors. Finally, we identify a glycinergic cell that is a potential muscarinic receptor-bearing target of GABA(B)-modulated acetylcholine release. This target is the DAPI-3 cell. We propose, based on these data, a modification of the Neal and Cunningham model in which GABA(B) receptors are on starburst, not glycinergic amacrine cells.

Amacrine Cells↗

PubMed2005

Ganglion cell densities in normal and dark-reared turtle retinas.

In dark-reared, neonatal turtle retinas, ganglion cell receptive fields and dendritic trees grow faster than normal. As a result, their areas may become, on average, up to twice as large as in control retinas. This raises the question of whether the coverage factor of dark-reared ganglion cells is larger than normal. Alternatively, dark rearing may lead to smaller-than-normal cell densities by accelerating apoptosis. To test these alternatives, we investigated the effect of light deprivation on densities and soma sizes of turtle retinal ganglion cells. For this purpose, we marked these cells using retrograde labeling of fixed turtle retinas with DiI (1,1'-dioctadecyl-3,3,3',3'-tetramethylindocarbocyanine perchlorate). Control turtles were maintained in a regular 12-h light/dark cycle from hatching until 4 weeks of age, whereas dark-reared turtles were maintained in total darkness for the same period. Ganglion cells in the control and dark-reared retinas were found to be similar in density and soma sizes. These results show that the mean coverage factor of turtle dark-reared ganglion cells is larger than normal.

Animals↗

PubMed2005

Measurement of rate of expansion in the perception of radial motion.

Optic flow generated by rigid surface patches can be decomposed into a small number of elementary motion types. In these experiments, we show that the human visual system can evaluate expansion, one of these motion types, metrically. Moreover, we show that the discrimination of rates of expansion are spatially local. Because the estimation of the focus of expansion is somewhat imprecise, this locality sometimes produces predictable errors in the estimation of rate of expansion. One can make predictions like this with a model adapted from one previously developed for angular-velocity discrimination.

Discrimination, Psychological↗

PubMed2005

Parametric decomposition of optic flow by humans.

Ego motion and natural motions in the world generate complex optic flows in the retina. These optic flows, if produced by rigid surface patches, can be decomposed into four components, including rotation and expansion. We showed previously that humans can precisely estimate parameters of these components, such as the angular velocity of a rotational motion and the rate of expansion of a radial motion. However, natural optic flows mostly display motions containing a combination of more than one of these components. Here, we report that when a pure motion (e.g., rotation) is combined with its orthogonal component (e.g., expansion), no bias is found in the estimate of the component parameters. This suggests that the visual system can decompose complex motions. However, this decomposition is such that the presence of the orthogonal component increases the discrimination threshold for the original component. We propose a model for how the brain decomposes the optic flow into its elementary components. The model accounts for how errors in the estimate of local-velocity vectors affect the decomposition, producing the increase of discrimination thresholds.

Brain↗

PubMed2003

Power spectra and distribution of contrasts of natural images from different habitats.

Some theories for visual receptive fields postulate that they depend on the image statistics of the natural habitat. Consequently, different habitats may lead to different receptive fields. We thus decided to study how some of the most relevant statistics vary across habitats. In particular, atmospheric and underwater habitats were compared. For these habitats, we looked at two measures of the power spectrum and one of the distributions of contrasts. From power spectra, we analyzed the log-log slope of the fall and the degree of isotropy. From the distribution of contrasts, we analyzed the fall in a semi-log scale. Past studies found that the spatial power spectra of natural atmospheric images fall linearly in logarithmic axes with a slope of about -2 and that their distribution of contrasts shows an approximate linear fall in semi-logarithmic axes. Here, we show that the power spectrum of underwater images have statistically significantly steeper slopes ( approximately -2.5 in log-log axes) than atmospheric images. The vast majority of power spectra are non-isotropic, but their degree of anisotropy is extremely low, especially in atmospheric images. There are also statistical differences across habitats for the distribution of contrasts, with it falling faster for underwater images than for atmospheric ones. We will argue that these differences are due to the optical properties of water and that the differences have relevance for theories of visual receptive fields. These theories would predict larger receptive fields for aquatic animals compared to land animals.

Contrast Sensitivity↗

PubMed2003

Local computation of angular velocity in rotational visual motion.

Retinal images evolve continuously over time owing to self-motions and to movements in the world. Such an evolving image, also known as optic flow, if arising from natural scenes can be locally decomposed in a Bayesian manner into several elementary components, including translation, expansion, and rotation. To take advantage of this decomposition, the brain has neurons tuned to these types of motions. However, these neurons typically have large receptive fields, often spanning tens of degrees of visual angle. Can neurons such as these compute elementary optic-flow components sufficiently locally to achieve a reasonable decomposition? We show that human discrimination of angular velocity is local. Local discrimination of angular velocity requires an accurate estimation of the center of rotation within the optic-flow field. Inaccuracies in estimating the center of rotation result in a predictable systematic error when one is estimating local angular velocity. Our results show that humans make the predicted errors. We discuss how the brain might estimate the elementary components of the optic flow locally by using large receptive fields.

Brain↗

PubMed2003

Sensory adaptation as Kalman filtering: theory and illustration with contrast adaptation.

Sensory adaptation allows biological systems to adjust to variations in the environment. A recent theoretical work postulated that the goal of adaptation is to minimize errors in the performance of particular tasks. The proposed minimization was Bayesian and required prior knowledge of the environment and of the limitations of the mechanisms processing the information. One problem with that formulation is that the environment changes in time and the theory did not specify how to know what the current state of the environment is. Here, we extend that theory to estimate optimally the environmental state from the temporal stream of responses. We show that such optimal estimation is a generalized form of Kalman filtering. An application of this new Kalman-filtering framework is worked out for retinal contrast adaptation. It is shown that this application can account for surprising features of the data. For example, it accounts for the differences in responses to increases and decreases of mean contrasts in the environment. In addition, it accounts for the two-phase decay of contrast gain when the mean contrast in the environment rises suddenly. The success of this and related theories suggest that sensory adaptation is a form of constrained biological optimization.

Adaptation, Physiological↗

PubMed2002

Measurement of angular velocity in the perception of rotation.

Humans are sensitive to the parameters of translational motion, namely, direction and speed. At the same time, people have special mechanisms to deal with more complex motions, such as rotations and expansions. One wonders whether people may also be sensitive to the parameters of these complex motions. Here, we report on a series of experiments that explore whether human subjects can use angular velocity to evaluate how fast a rotational motion is. In four experiments, subjects were required to perform a task of speed-of-rotation discrimination by comparing two annuli of different radii in a temporal 2AFC paradigm. Results showed that humans could rely on a sensitive measurement of angular velocity to perform this discrimination task. This was especially true when the quality of the rotational signal was high (given by the number of dots composing the annulus). When the signal quality decreased, a bias towards linear velocity of 5-80% appeared, suggesting the existence of separate mechanisms for angular and linear velocity. This bias was independent from the reference radius. Finally, we asked whether the measurement of angular velocity required a rigid rotation, that is, whether the visual system makes only one global estimate of angular velocity. For this purpose, a random-dot disk was built such that all the dots were rotating with the same tangential speed, irrespectively of radius. Results showed that subjects do not estimate a unique global angular velocity, but that they perceive a non-rigid disk, with angular velocity falling inversely proportionally with radius.

Humans↗

PubMed2002

Temporal coherence in visual rotation.

The brain can predict and estimate motion based on visual translation. This paper addresses whether the visual system also has a specialized mechanism of temporal coherence for rotational motion. To do this, we measured the perceived mislocation of a rotating dot at the time of its luminance transition. Results show that subjects mislocate the dot consistently with its circular motion rather than with translational temporal coherence. We propose a model to explain these results based on a combination of an error in a location-estimation task and on the brain assuming rotational motions.

Humans↗

PubMed2002

A Bayesian framework for sensory adaptation.

Adaptation allows biological sensory systems to adjust to variations in the environment and thus to deal better with them. In this article, we propose a general framework of sensory adaptation. The underlying principle of this framework is the setting of internal parameters of the system such that certain prespecified tasks can be performed optimally. Because sensorial inputs vary probabilistically with time and biological mechanisms have noise, the tasks could be performed incorrectly. We postulate that the goal of adaptation is to minimize the number of task errors. This minimization requires prior knowledge of the environment and of the limitations of the mechanisms processing the information. Because these processes are probabilistic, we formulate the minimization with a Bayesian approach. Application of this Bayesian framework to the retina is successful in accounting for a host of experimental findings.

Adaptation, Physiological↗

PubMedDate not supplied

Stimulus-dependent correlated firing in directionally selective retinal ganglion cells.

Synchronous spiking has been postulated to be a meta-signal in visual cortex and other CNS loci that tags neuronal spike responses to a single entity. In retina, however, synchronized spikes have been postulated to arise via mechanisms that would largely preclude their carrying such a code. One such mechanism is gap junction coupling, in which synchronous spikes would be a by-product of lateral signal sharing. Synchronous spikes have also been postulated to arise from common-source inputs to retinal ganglion cells having overlapping receptive fields, and thus code for stimulus location in the overlap area. On-Off directionally selective ganglion cells of the rabbit retina exhibit a highly precise tiling pattern in which gap junction coupling occurs between some neighboring, same-preferred-direction cells. Depending on how correlated spikes arise, and for what purpose, one could postulate that synchronized spikes in this system (1) always arise in some subset of same-direction cells because of gap junctions, but never in non-same-preferred-directional cells; (2) never arise in same-directional cells because their receptive fields do not overlap, but arise only in different-directional cells whose receptive fields overlap, as a code for location in the overlap region; or (3) arise in a stimulus-dependent manner for both same- and different-preferred-direction cells for a function similar to that postulated for neurons in visual cortex. Simultaneous, extracellular recordings were obtained from neighboring On-Off directionally selective (DS) ganglion cells having the same and different preferred directions in an isolated rabbit retinal preparation. Stimulation by large flashing spots elicited responses from DS ganglion-cell pairs that typically showed little synchronous firing. Movement of extended bars, however, often produced synchronous spikes in cells having similar or orthogonal preferred directions. Surprisingly, correlated firing could occur for the opposite contrast polarity edges of moving stimuli when the leading edge of a sweeping bar excited the receptive field of one cell as its trailing edge stimulated another. Pharmacological manipulations showed that the spike synchronization is enhanced by excitatory cholinergic amacrine-cell inputs, and reduced by inhibitory GABAergic inputs, in a motion-specific manner. One possible interpretation is that this synchronous firing could be a signal to higher centers that the outputs of the two DS ganglion cells should be "bound" together as responding to a contour of a common object.

Animals↗