scholarly journals The direction-selective contrast response of area 18 neurons is different for first- and second-order motion

2005 ◽  
Vol 22 (1) ◽  
pp. 87-99 ◽  
Author(s):  
TIMOTHY LEDGEWAY ◽  
CHANG'AN ZHAN ◽  
AARON P. JOHNSON ◽  
YUNING SONG ◽  
CURTIS L. BAKER

Cortical neurons selective for the direction of motion often exhibit some limited response to motion in their nonpreferred directions. Here we examine the dependence of neuronal direction selectivity on stimulus contrast, both for first-order (luminance-modulated, sine-wave grating) and second-order (contrast-modulated envelope) stimuli. We measured responses from single neurons in area 18 of cat visual cortex to both kinds of moving stimuli over a wide range of contrasts (1.25–80%). Direction-selective contrast response functions (CRFs) were calculated as the preferred-minus-null difference in average firing frequency as a function of contrast. We also applied receiver operating characteristic analysis to our CRF data to obtain neurometric functions characterizing the potential ability of each neuron to discriminate motion direction at each contrast level tested. CRFs for sine-wave gratings were usually monotonic; however, a substantial minority of neurons (35%) exhibited nonmonotonic CRFs (such that the degree of direction selectivity decreased with increasing contrast). The underlying preferred and nonpreferred direction CRFs were diverse, often having different shapes in a given neuron. Neurometric functions for direction discrimination showed a similar degree of heterogeneity, including instances of nonmonotonicity. For contrast-modulated stimuli, however, CRFs for either carrier or envelope contrast were always monotonic. In a given neuron, neurometric thresholds were typically much higher for second- than for first-order stimuli. These results demonstrate that the degree of a cell's direction selectivity depends on the contrast at which it is measured, and therefore is not a characteristic parameter of a neuron. In general, contrast response functions for first-order stimuli were very heterogeneous in shape and sensitivity, while those for second-order stimuli showed less sensitivity and were quite stereotyped in shape.

2005 ◽  
Vol 94 (2) ◽  
pp. 1336-1345 ◽  
Author(s):  
Bartlett D. Moore ◽  
Henry J. Alitto ◽  
W. Martin Usrey

The activity of neurons in primary visual cortex is influenced by the orientation, contrast, and temporal frequency of a visual stimulus. This raises the question of how these stimulus properties interact to shape neuronal responses. While past studies have shown that the bandwidth of orientation tuning is invariant to stimulus contrast, the influence of temporal frequency on orientation-tuning bandwidth is unknown. Here, we investigate the influence of temporal frequency on orientation tuning and direction selectivity in area 17 of ferret visual cortex. For both simple cells and complex cells, measures of orientation-tuning bandwidth (half-width at half-maximum response) are ∼20–25° across a wide range of temporal frequencies. Thus cortical neurons display temporal-frequency invariant orientation tuning. In contrast, direction selectivity is typically reduced, and occasionally reverses, at nonpreferred temporal frequencies. These results show that the mechanisms contributing to the generation of orientation tuning and direction selectivity are differentially affected by the temporal frequency of a visual stimulus and support the notion that stability of orientation tuning is an important aspect of visual processing.


1980 ◽  
Vol 47 (1) ◽  
pp. 75-81 ◽  
Author(s):  
R. T. Shield

When a mechanical system has a potential energy, it is a simple matter to show that if the generalized force corresponding to a coordinate p is known to first order in p for a range of the other coordinates of the system, then the other generalized forces can be found immediately to second order in p, without requiring a second-order analysis of the system. By this method the second-order change in the axial force when a finitely extended elastic cylinder is twisted is found from the first-order value of the twisting moment. Numerical results for a realistic form of the strain-energy function for an incompressible material suggest that the second-order expression for the axial force is very accurate for a wide range of twist for circular cylinders of rubber-like materials extended 100 percent or more.


1985 ◽  
Vol 54 (3) ◽  
pp. 651-667 ◽  
Author(s):  
I. Ohzawa ◽  
G. Sclar ◽  
R. D. Freeman

We have examined the idea that the adaptation of cortical neurons to local contrast levels in a visual stimulus is functionally advantageous. Specifically, cortical cells may have large differential contrast sensitivity as a result of adjustments that center a limited response range around a mean level of contrast. To evaluate this notion, we measured contrast-response functions of cells in striate cortex while systematically adapting them to different contrast levels of stimulus gratings. For the majority of cortical neurons tested, the results of this basic experiment show that contrast-response functions shift laterally along a log-contrast axis so that response functions match mean contrast levels in the stimulus. This implies a contrast-dependent change in the gain of the cell's contrast-response relationship. We define this process as contrast gain control. The degree to which this contrast adjustment occurs varies considerably from cell to cell. There are no obvious differences regarding cell type (simple vs. complex) or laminar distribution. Contrast gain control is almost certainly a cortical function, since lateral geniculate cells and fibers exhibit only minimal effects. Tests presented in the accompanying paper (37) provide additional evidence on the cortical origin of the process. In another series of experiments, the effect of contrast adaptation on physiological estimates of contrast sensitivity was evaluated. Sustained adaptation to contrast levels as low as 3% was capable of nearly doubling the thresholds of most of the cells tested. Adaptation may therefore be an important factor in determinations of the contrast sensitivity of cortical neurons. We tested the spatial extent of the mechanisms responsible for these gain-control effects by attempting to adapt cells using both a large grating and a grating patch limited to that portion of a cell's receptive field from which excitatory discharges could be elicited directly (the central discharge region). Adaptation was found to be an exclusive property of the central region. This held even in the case of hypercomplex cells, which received strong influences from surrounding regions of the visual field. Finally, we measured the time course of contrast adaptation. We found the process to be rather slow, with a mean time constant of approximately 6 s. Once again, there was considerable variability in this value from cell to cell.


1997 ◽  
Vol 78 (4) ◽  
pp. 2034-2047 ◽  
Author(s):  
Hiroko M. Sakai ◽  
Hildred Machuca ◽  
Michael J. Korenberg ◽  
Ken-Ichi Naka

Sakai, Hiroko M., Hildred Machuca, Michael J. Korenberg, and Ken-Ichi Naka. Processing of color- and noncolor-coded signals in the gourami retina. III. Ganglion cells. J. Neurophysiol. 78: 2034–2047, 1997. The dynamics of intracellular responses from ganglion cells, as well as that of spike discharges, were studied with the stimulus regimens and analytic procedures identical to those used to study the dynamics of the responses from horizontal and amacrine cells ( Sakai et al. 1997a , b ). The stimuli used were large fields of red and green light given as a pulsatile input or modulation about a mean luminance by a white-noise signal. Spike discharges evoked by a white-noise stimulus were analyzed in exactly the same manner as that used for analysis of analog responses. The canonical nature of kernels allowed us to correlate the first- and second-order components in a spike train with those of the intracellular responses from horizontal, amacrine, and ganglion cells. Both red and green stimuli given alone in darkness produced noncolor-coded responses from all ganglion cells. In the case of some cells, steady red illumination changed the polarity or waveform of the response to green light. Color-coded ganglions responded only to simultaneous color contrast. Nonlinearities recovered from intracellular responses, and spike discharges were similar to those found in responses from amacrine cells and were of two types, one characteristic of the C amacrine cells and the other characteristic of the N amacrine cells. The first-order kernels of most ganglion cells could be divided into two basic types, biphasic and triphasic. The combination of kernels of these two basic types with different polarities can produce a wide range of responses. Addition of two types of second-order nonlinearity could render color coding in this relatively simple retina as an extremely complex process. Color information appeared to be represented by the polarity, as well as the waveform, of the first-order kernel. The response dynamics is a means of transmission of color-coded information. Second-order components carry information about changes around a mean luminance regardless of the color of an input. Some spike discharges produced a well-defined cross-kernel between red and green inputs to show that a particular time sequence of red and green stimuli was detected by the retinal neuron network. The similarity between signatures of second-order kernels for both amacrine and ganglion cells indicates that signals undergo a minimal transformation in the temporal domain when they are transmitted from amacrine to ganglion cells and then transformed into a spike train. Under our experimental conditions, a single spike train carried simultaneously information about red and green inputs, as well as about linear and nonlinear components. In addition, the spike train also carries a cross-talk component. A spike train is a carrier of multiple signals. Conversely, many types of information in a stimulus are independently encoded into a spike train.


1985 ◽  
Vol 54 (3) ◽  
pp. 668-675 ◽  
Author(s):  
G. Sclar ◽  
I. Ohzawa ◽  
R. D. Freeman

We have studied the effects of contrast adaptation on cortical cells from 4- and 6-wk-old kittens (49 and 47 cells, respectively) using sine-wave grating stimuli. We wished to know if the effects of adaptation to different contrast levels are more extensive than those in adult animals. Our experiments involved adapting cells to different contrasts (3.1, 12.5, and 50%) while concurrently measuring their contrast-response functions at each of these different levels. We found qualitatively that the effects of adaptation in the kitten are similar to those we have previously documented in adult animals (19). Contrast-response functions are laterally shifted along the log-contrast axis, effectively matching the response range of the cells to prevailing contrast levels. The degree to which this occurred varied from cell to cell. The average degree to which cells showed these effects, as assessed both qualitatively and quantitatively, was greater for kittens than for adult cats, and greater for 4-wk-old kittens than for those aged 6 wk. This suggests that susceptibility to adaptation varies as a function of age. Additional studies were undertaken with the intent of localizing these adaptive effects. First, lateral geniculate cells and fibers (n = 23) were studied with our standard protocol, and second, we investigated the degree to which the effects of adaptation of cortical cells transferred interocularly.(ABSTRACT TRUNCATED AT 250 WORDS)


Materials ◽  
2021 ◽  
Vol 14 (18) ◽  
pp. 5384
Author(s):  
Mohamad Al Nashar ◽  
Alok Sutradhar

Hierarchical lattices are structures composed of self-similar or dissimilar architected metamaterials that span multiple length scales. Hierarchical lattices have superior and tunable properties when compared to conventional lattices, and thus, open the door for a wide range of material property manipulation and optimization. Using finite element analysis, we investigate the energy absorption capabilities of 3D hierarchical lattices for various unit cells under low strain rates and loads. In this study, we use fused deposition modeling (FDM) 3D printing to fabricate a dog bone specimen and extract the mechanical properties of thermoplastic polyurethane (TPU) 85A with a hundred percent infill printed along the direction of tensile loading. With the numerical results, we observed that the energy absorption performance of the octet lattice can be enhanced four to five times by introducing a hierarchy in the structure. Conventional energy absorption structures such as foams and lattices have demonstrated their effectiveness and strengths; this research aims at expanding the design domain of energy absorption structures by exploiting 3D hierarchical lattices. The result of introducing a hierarchy to a lattice on the energy absorption performance is investigated by varying the hierarchical order from a first-order octet to a second-order octet. In addition, the effect of relative density on the energy absorption is isolated by creating a comparison between a first-order octet lattice with an equivalent relative density as a second-order octet lattice. The compression behaviors for the second order octet, dodecahedron, and truncated octahedron are studied. The effect of changing the cross-sectional geometry of the lattice members with respect to the energy absorption performance is investigated. Changing the orientation of the second-order cells from 0 to 45 degrees has a considerable impact on the force–displacement curve, providing a 20% increase in energy absorption for the second-order octet. Analytical solutions of the effective elasticity modulus for the first- and second-order octet lattices are compared to validate the simulations. The findings of this paper and the provided understanding will aid future works in lattice design optimization for energy absorption.


1997 ◽  
Vol 36 (04/05) ◽  
pp. 315-318 ◽  
Author(s):  
K. Momose ◽  
K. Komiya ◽  
A. Uchiyama

Abstract:The relationship between chromatically modulated stimuli and visual evoked potentials (VEPs) was considered. VEPs of normal subjects elicited by chromatically modulated stimuli were measured under several color adaptations, and their binary kernels were estimated. Up to the second-order, binary kernels obtained from VEPs were so characteristic that the VEP-chromatic modulation system showed second-order nonlinearity. First-order binary kernels depended on the color of the stimulus and adaptation, whereas second-order kernels showed almost no difference. This result indicates that the waveforms of first-order binary kernels reflect perceived color (hue). This supports the suggestion that kernels of VEPs include color responses, and could be used as a probe with which to examine the color visual system.


2017 ◽  
Vol 9 (3) ◽  
pp. 17-30
Author(s):  
Kelly James Clark

In Branden Thornhill-Miller and Peter Millican’s challenging and provocative essay, we hear a considerably longer, more scholarly and less melodic rendition of John Lennon’s catchy tune—without religion, or at least without first-order supernaturalisms (the kinds of religion we find in the world), there’d be significantly less intra-group violence. First-order supernaturalist beliefs, as defined by Thornhill-Miller and Peter Millican (hereafter M&M), are “beliefs that claim unique authority for some particular religious tradition in preference to all others” (3). According to M&M, first-order supernaturalist beliefs are exclusivist, dogmatic, empirically unsupported, and irrational. Moreover, again according to M&M, we have perfectly natural explanations of the causes that underlie such beliefs (they seem to conceive of such natural explanations as debunking explanations). They then make a case for second-order supernaturalism, “which maintains that the universe in general, and the religious sensitivities of humanity in particular, have been formed by supernatural powers working through natural processes” (3). Second-order supernaturalism is a kind of theism, more closely akin to deism than, say, Christianity or Buddhism. It is, as such, universal (according to contemporary psychology of religion), empirically supported (according to philosophy in the form of the Fine-Tuning Argument), and beneficial (and so justified pragmatically). With respect to its pragmatic value, second-order supernaturalism, according to M&M, gets the good(s) of religion (cooperation, trust, etc) without its bad(s) (conflict and violence). Second-order supernaturalism is thus rational (and possibly true) and inconducive to violence. In this paper, I will examine just one small but important part of M&M’s argument: the claim that (first-order) religion is a primary motivator of violence and that its elimination would eliminate or curtail a great deal of violence in the world. Imagine, they say, no religion, too.Janusz Salamon offers a friendly extension or clarification of M&M’s second-order theism, one that I think, with emendations, has promise. He argues that the core of first-order religions, the belief that Ultimate Reality is the Ultimate Good (agatheism), is rational (agreeing that their particular claims are not) and, if widely conceded and endorsed by adherents of first-order religions, would reduce conflict in the world.While I favor the virtue of intellectual humility endorsed in both papers, I will argue contra M&M that (a) belief in first-order religion is not a primary motivator of conflict and violence (and so eliminating first-order religion won’t reduce violence). Second, partly contra Salamon, who I think is half right (but not half wrong), I will argue that (b) the religious resources for compassion can and should come from within both the particular (often exclusivist) and the universal (agatheistic) aspects of religious beliefs. Finally, I will argue that (c) both are guilty, as I am, of the philosopher’s obsession with belief. 


2009 ◽  
Vol 74 (1) ◽  
pp. 43-55 ◽  
Author(s):  
Dennis N. Kevill ◽  
Byoung-Chun Park ◽  
Jin Burm Kyong

The kinetics of nucleophilic substitution reactions of 1-(phenoxycarbonyl)pyridinium ions, prepared with the essentially non-nucleophilic/non-basic fluoroborate as the counterion, have been studied using up to 1.60 M methanol in acetonitrile as solvent and under solvolytic conditions in 2,2,2-trifluoroethan-1-ol (TFE) and its mixtures with water. Under the non- solvolytic conditions, the parent and three pyridine-ring-substituted derivatives were studied. Both second-order (first-order in methanol) and third-order (second-order in methanol) kinetic contributions were observed. In the solvolysis studies, since solvent ionizing power values were almost constant over the range of aqueous TFE studied, a Grunwald–Winstein equation treatment of the specific rates of solvolysis for the parent and the 4-methoxy derivative could be carried out in terms of variations in solvent nucleophilicity, and an appreciable sensitivity to changes in solvent nucleophilicity was found.


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