Biomedical subjects
Publications and source records attributed to A W Przybyszewski.
Anatomical studies of the visual cortex have identified massive back-projecting pathways. Theoretical studies suggest how such pathways may play important roles in vision by mediating 'top-down' processing, in which information from a relatively high level is fed back to early visual stages.
Enroth-Cugell and Robson (1966) first proposed a classification of retinal ganglion cells into X cells, which exhibit approximate linear spatial summation and largely sustained responses, and Y cells, which exhibit nonlinearities and transient responses. Gaudiano (1992a, 1992b, 1994) has suggested that the dominant characteristics of both X and Y cells can be simulated with a single model simply by changing receptive field profiles to match those of the anatomical counterparts of X and Y cells. He also proposed that a significant component of the spatial nonlinearities observed in Y (and sometimes X) cells can result from photoreceptor nonlinearities coupled with push-pull bipolar connections. Specifically, an asymmetry was predicted in the ganglion cell response to rectangular gratings presented at different locations in the receptive field under two conditions: introduction/withdrawal (on-off) or contrast reversal. When measuring the response to these patterns as a function of spatial phase, the standard difference-of-Gaussians model predicts symmetrical responses about the receptive field center, while the push-pull model predicts slight but significant asymmetry in the on-off case only. To test this hypothesis, we have recorded ganglion cell responses from the optic tract fibers of anesthetized cat. The mean and standard deviations of responses to on-off and contrast-reversed patterns were compared. We found that all but one of the cells that yielded statistically significant data confirmed the hypothesis. These results largely support the theoretical prediction.
We recorded intracellular responses from cat retinal ganglion cells to sinusoidal flickering lights, and compared the response dynamics with a theoretical model based on coupled nonlinear oscillators. Flicker responses for several different spot sizes were separated in a "smooth" generator (G) potential and corresponding spike trains. We have previously shown that the G-potential reveals complex, stimulus-dependent, oscillatory behavior in response to sinusoidally flickering lights. Such behavior could be simulated by a modified van der Pol oscillator. In this paper, we extend the model to account for spike generation as well, by including extended Hodgkin-Huxley equations describing local membrane properties. We quantified spike responses by several parameters describing the mean and standard deviation of spike burst duration, timing (phase shift) of bursts, and the number of spikes in a burst. The dependence of these response parameters on stimulus frequency and spot size could be reproduced in great detail by coupling the van der Pol oscillator and Hodgkin-Huxley equations. The model mimics many experimentally observed response patterns, including non-phase-locked irregular oscillations. Our findings suggest that the information in the ganglion cell spike train reflects both intraretinal processing, simulated by the van der Pol oscillator, and local membrane properties described by Hodgkin-Huxley equations. The interplay between these complex processes can be simulated by changing the coupling coefficients between the two oscillators. Our simulations therefore show that irregularities in spike trains, which normally are considered to be noise, may be interpreted as complex oscillations that might carry information.
Intracellularly recorded light-responses of X-type ganglion cells in the cat retina were separated, with the help of a wavelet method, into "slow" membrane ("G")-potentials and the corresponding spike trains. In response to sinusoidally modulated high intensity light spots with different sizes and frequencies, X-type ganglion cells show both oscillations correlated with the stimulus frequency and other, faster, oscillations that were not always locked to the stimulus. A forced van der Pol oscillator model with stimulus-dependent coefficients proved to describe the empirical findings quite well. A linearity-coefficient of the equations indicates strong nonlinearity at a temporal frequency of 8 Hz with spot sizes on the order of 0.5-0.7 deg and decreasing nonlinearity at lower temporal frequencies or smaller spot sizes, while the faster oscillations become more prominent. We could not determine whether the oscillations are intrinsic to the cell-membrane or generated by (or in interaction with) the preganglionic retinal meshwork. The results show that X-cell spike-trains can contain oscillations that are not phase-locked to the stimulus and that are therefore virtually invisible after stimulus synchronous averaging. It is not likely that these retinal oscillations directly induce the well described oscillations in cat visual cortex, since they usually fall in a different frequency range.
To investigate the sites of light adaptation processes in the mammalian distal retina, we studied the lateral spread of adaptation signals in cone-driven cat horizontal (H-) cell responses. The size of the adaptation pool is compared to the receptive field for H-cell responses. H-cell activity was recorded intracellularly in the optically intact, in vivo eye. It is demonstrated that light adaptation as measured in H-cells is not a strictly local process. Background light falling outside a central test region effectively modulates the responses to a small test light, flashed on the receptive field center. The integration area for adaptation signals was quantitatively compared to the H-cell receptive field size by measuring the desensitizing effect of background light on the responses to a small centered test spot, as a function of background spot size. The area-adaptation function is comparable to the area-response function but has a slightly smaller length constant. Light adaptation in H-cell responses, therefore, reveals spread of adaptation over a large distance and is probably mediated through lateral interactions in the H-cell network rather than in the cones.
This paper discusses a simple application of the wavelet transformation to analyse nerve cell impulse patterns. The action potentials converted into delta, or Dirac, functions were convoluted in the time domain with a modified Gauss (the negative of the second derivative of Gauss) function, varying in width between 0.6 and 384 ms. The width of the Gauss function was varied in 640 steps. Some parts of the transformation were extended, analysed and averaged in the frequency domain to explore oscillatory components of the impulse pattern. The sequences of action potentials of retinal ganglion cells evoked by short flashes are taken as examples. The present analysis demonstrate some properties of mathematical "microscopic" application to transient responses of the central nervous system (CNS), whereby the degree of magnification (steps of transformation) was varied.
Eyeball deformation in total darkness leads to an activation of on-center ganglion cells and an inhibition of off-center ganglion cells. After "deformation off" most on-center ganglion cell activity decreased slowly to the normal spontaneous dark level, while in off-center ganglion cells some returned according to an exponential function to normal dark activity, while others had a transient postinhibitory activation period. In general, the response type of latency class I and latency class II neurons was the same. Dark adaptation of 30-45 min duration only changed this neuronal response pattern slightly, if at all. A detailed statistical analysis is provided for the four classes of retinal ganglion cells recorded: latency class I on-center and off-center neurons and latency class II on-center and off-center neurons. The missing effects of dark adaptation on neuronal responses evoked by eyeball deformation are explained by three possible models. The more plausible one assumes that horizontal cells are depolarized by retinal stretch. Their interaction with cone on-bipolars or cone off-bipolars is fairly independent of photoreceptor adaptation or transmitter release at the cone pedicles and is still effective when all molecular receptor sites at cone/bipolar cell synapses are occupied during scotopic states of dark adaptation. In psychophysical experiments (two subjects), as in the neuronal responses, we also could not find any indication that the "pressure phosphenes" evoked by lateral eyeball indentation are altered during dark adaptation.
By means of microelectrodes, the activity of single neurons (on-center, off-center ganglion cells, latency class I and class II neurons) was recorded from the optic tract of anesthetized cats. Eyeball deformation in total darkness led fairly consistently to an activation of the on-center ganglion cells, while off-center ganglion cells were inhibited. The latency and strength of this activation or inhibition seemed to be mainly dependent on the strength of eyeball indentation and the location of the neurons relative to the point of eyeball indentation. Some on-center neurons (mostly latency class I) also exhibited a short activation at "deformation off". For comparison, the responses of retinal ganglion cells to eyeball deformation in a hydrostatically open system and to a sudden increase in the intraocular pressure (closed system) are described. The neurophysiological data are explained by the assumption that eyeball indentation leads to a nonuniform tangential stretch of the retina, which exerts a locally variable depolarization of horizontal cells. This horizontal cell depolarization leads either directly or via a feedback loop through cone pedicles to a depolarization of on-bipolars and a hyperpolarization of off-bipolars. These effects determine in turn the responses seen at the ganglion cell level. It is emphasized that eyeball deformation can be used as an independent tool in transmitter studies of the retina.
(a) When a subject indents one of his eyeballs in total darkness, he immediately perceives light extending slowly across the whole visual field of the indented eye. The appearance and the time course of these pressure or deformation phosphenes are described. (b) With simultaneous binocular indentation of the eyeballs a flickering patterned phosphene is observed. (c) A short history of the research on pressure phosphenes and its consequences for the theories of vision is presented. (d) Purkynĕ's observations of monocular deformation phosphenes are described. He repeatedly noted patterned light structures, which most observers only perceive with simultaneous binocular eyeball deformation. It is suggested that Purkynĕ's deviating observations were caused by amblyopia of one eye. (e) The neurophysiological basis of the monocular pressure phosphenes was investigated by means of microelectrode recordings from single optic tract fibers. The activity of single retinal ganglion cells (on-center, off-center neurons, latency class I [Y-neurons] or latency class II [X-neurons]), was recorded in anaesthetized cats. Eyeball deformation in total darkness led to an activation of the on-center ganglion cells, while the off-center ganglion cells were inhibited. The latency and strength of this activation or inhibition varied considerably between different neurons, but were fairly constant in the same neuron when the eyeball indentation was repeated after a pause of 1-3 min. The latency and strength of neuronal activation or inhibition seemed to be dependent mainly upon the neuron location relative to the point of eyeball indentation. Some on-center neurons also exhibited a short activation at "deformation off". (f) The antagonistic response type of on-center and off-center ganglion cells was also observed when the eyeball was deformed as a hydrostatic open system and the intraocular pressure was kept at 25 mm Hg basic pressure. (g) Dark adaptation up to 45 min affected the deformation responses of retinal neurons only to a small degree, if at all. This corresponds to the observation that deformation phosphenes in a human observer changed little during the course of dark adaptation. (h) We assume that the activation of on-center and inhibition of off-center ganglion cells by eyeball deformation are caused by retinal stretching, which also leads to horizontal cell stretch. Stretching the horizontal cell membrane probably generates an increase in membrane sodium conductivity and a depolarization of the membrane potential. This depolarization of the horizontal cell membrane potential is transmitted either directly or indirectly (via receptor synapses) from the horizontal to the bipolar cells.(ABSTRACT TRUNCATED AT 400 WORDS)
Recurrent projections comprise a universal feature of cerebral organization. Here, we show that the corticofugal projections from the striate cortex (VI) to the lateral geniculate nucleus (LGN) robustly and multiplicatively enhance the responses of parvocellular neurons, stimulated by gratings restricted to the classical receptive field and modulated in luminance, by over two-fold in a contrast-independent manner at all but the lowest contrasts. In the equiluminant plane, wherein stimuli are modulated in chromaticity with luminance held constant, such enhancement is strongly contrast dependent. These projections also robustly enhance the responses of magnocellular neurons but contrast independently only at high contrasts. Thus, these results have broad functional significance at both network and neuronal levels by providing the experimental basis and quantitative constraints for a wide range of models on recurrent projections and the control of contrast gain.