Contrast-invariant orientation tuning in cat visual cortex: thalamocortical input tuning and correlation-based intracortical connectivity

The origin of orientation selectivity in visual cortical responses is a central problem for understanding cerebral cortical circuitry. In cats, many experiments suggest that orientation selectivity arises from the arrangement of lateral geniculate nucleus (LGN) afferents to layer 4 simple cells. However, this explanation is not sufficient to account for the contrast invariance of orientation tuning. To understand contrast invariance, we first characterize the input to cat simple cells generated by the oriented arrangement of LGN afferents. We demonstrate that it has two components: a spatial-phase-specific component (i.e., one that depends on receptive field spatial phase), which is tuned for orientation, and a phase-nonspecific component, which is untuned. Both components grow with contrast. Second, we show that a correlation-based intracortical circuit, in which connectivity between cell pairs is determined by the correlation of their LGN inputs, is sufficient to achieve well tuned, contrast-invariant orientation tuning. This circuit generates both spatially opponent, "antiphase" inhibition ("push-pull"), and spatially matched, "same-phase" excitation. The inhibition, if sufficiently strong, suppresses the untuned input component and sharpens responses to the tuned component at all contrasts. The excitation amplifies tuned responses. This circuit agrees with experimental evidence showing spatial opponency between, and similar orientation tuning of, the excitatory and inhibitory inputs received by a simple cell. Orientation tuning is primarily input driven, accounting for the observed invariance of tuning width after removal of intracortical synaptic input, as well as for the dependence of orientation tuning on stimulus spatial frequency. The model differs from previous push-pull models in requiring dominant rather than balanced inhibition and in predicting that a population of layer 4 inhibitory neurons should respond in a contrast-dependent manner to stimuli of all orientations, although their tuning width may be similar to that of excitatory neurons. The model demonstrates that fundamental response properties of cortical layer 4 can be explained by circuitry expected to develop under correlation-based rules of synaptic plasticity, and shows how such circuitry allows the cortex to distinguish stimulus intensity from stimulus form.     

Translation-invariant orientation tuning in visual "complex" cells could derive from intradendritic computations

first distinguished "simple" from "complex" cells in visual cortex and proposed a processing hierarchy in which rows of LGN cells are pooled to drive oriented simple cell subunits, which are pooled in turn to drive complex cells. Although parsimonious and highly influential, the pure hierarchical model has since been challenged by results indicating that many complex cells receive excitatory monosynaptic input from LGN cells or do not depend on simple cell input. Alternative accounts of complex cell orientation tuning remain scant, however, and the function of monosynaptic LGN contacts onto complex cell dendrites remains unknown. We have used a biophysically detailed compartmental model to investigate whether nonlinear integration of LGN synaptic inputs within the dendrites of individual pyramidal cells could contribute to complex-cell receptive field structure. We show that an isolated cortical neuron with "active" dendrites, driven only by excitatory inputs from overlapping ON- and OFF-center LGN subfields, can produce clear phase-invariant orientation tuning-a hallmark response characteristic of a complex cell. The tuning is shown to depend critically both on the spatial arrangement of LGN synaptic contacts across the complex cell dendritic tree, established by a Hebbian developmental principle, and on the physiological efficacy of excitatory voltage-dependent dendritic ion channels. We conclude that unoriented LGN inputs to a complex cell could contribute in a significant way to its orientation tuning, acting in concert with oriented inputs to the same cell provided by simple cells or other complex cells. As such, our model provides a novel, experimentally testable hypothesis regarding the basis of orientation tuning in the complex cell population, and more generally underscores the potential importance of nonlinear intradendritic subunit processing in cortical neurophysiology.     

Strobe rearing prevents the convergence of inputs with different response timings onto area 17 simple cells

Strobe rearing prevents the convergence of inputs with different response timings onto area 17 simple cells. J. Neurophysiol. 80: 3005-3020, 1998. The preceding paper showed that the loss of direction selectivity in simple cells induced by strobe rearing reflects the elimination of spatially ordered response timing differences across the receptive field that underlie spatiotemporal (S-T) inseparability. Here we addressed whether these changes reflected an elimination of certain timings or an alteration in how timings were associated in single cells. Timing in receptive fields was measured using stationary bars undergoing sinusoidal luminance modulation at different temporal frequencies (0.5-6 Hz). For each bar position, response phase versus temporal frequency data were fit by a line to obtain two measures: absolute phase and latency. In normal cats, many individual simple cells display a wide range of timings; in layer 4, the mean range for absolute phase and latency was 0.21 cycles and 39 ms, respectively. Strobe rearing compressed the mean timing ranges in single cells, to 0.08 cycles and 31 ms, respectively, and this compression accounted for the loss of inseparability. A similar compression was measured in layer 6 cells. In contrast, the range of timing values across the simple-cell population was relatively normal. Single cells merely sampled narrower than normal regions of the timing space. We sought to understand these cortical changes in terms of how inputs from the lateral geniculate nucleus (LGN) may have been affected by strobe rearing. In normal cats, a wide range of absolute phase and latency values exists among lagged and nonlagged LGN cells, and these thalamic timings account for most of the cortical timings. Also, S-T inseparability in many simple cells can be attributed to the convergence of lagged and/or nonlagged inputs. Strobe rearing did not change the sampling of lagged and nonlagged cells, and the geniculate timings continued to account for most of the cortical timings. However, strobe rearing virtually eliminated cortical receptive fields with mixed lagged and nonlagged timing, and it compressed the timing range in cells dominated by one or the other geniculate type. Thus strobe rearing did not eliminate certain timings in LGN or cortex, but prevented the convergence of different timings on single cells. To account for these results, we propose a developmental model in which strobe stimulation alters the correlational structure of inputs based on their response timing. Only inputs with similar timing become associated on single cortical cells, and this produces S-T separable receptive fields that lack the ability to confer a preferred direction of motion.     

Strobe rearing reduces direction selectivity in area 17 by altering spatiotemporal receptive-field structure

Strobe rearing reduces direction selectivity in area 17 by altering spatiotemporal receptive-field structure. J. Neurophysiol. 80: 2991-3004, 1998. Direction selectivity in simple cells of cat area 17 is linked to spatiotemporal (S-T) receptive-field structure. S-T inseparable receptive fields display gradients of response timing across the receptive field that confer a preferred direction of motion. Receptive fields that are not direction selective lack gradients; they are S-T separable, displaying uniform timing across the field. Here we further examine this link using a developmental paradigm that disrupts direction selectivity. Cats were reared from birth to 8 mo of age in 8-Hz stroboscopic illumination. Direction selectivity in simple cells was then measured using gratings drifting at different temporal frequencies (0.25-16 Hz). S-T structure was assessed using stationary bars presented at different receptive-field positions, with bar luminance being modulated sinusoidally at different temporal frequencies. For each cell, plots of response phase versus bar position were fit by lines to characterize S-T inseparability at each temporal frequency. Strobe rearing produced a profound loss of direction selectivity at all temporal frequencies; only 10% of cells were selective compared with 80% in normal cats. The few remaining directional cells were selective over a narrower than normal range of temporal frequencies and exhibited weaker than normal direction selectivity. Importantly, the directional loss was accompanied by a virtual elimination of S-T inseparability. Nearly all cells were S-T separable, like nondirectional cells in normal cats. The loss was clearest in layer 4. Normally, inseparability is greatest there, and it correlates well (r = 0.77) with direction selectivity; strobe rearing reduced inseparability and direction selectivity to very low values. The few remaining directional cells were inseparable. In layer 6 of normal cats, most direction-selective cells are only weakly inseparable, and there is no consistent relationship between the two measures. However, after strobe rearing, even the weak inseparability was eliminated along with direction selectivity. The correlated changes in S-T structure and direction selectivity were confirmed using conventional linear predictions of directional tuning based on responses to counterphasing bars and white noise stimuli. The developmental changes were permanent, being observed up to 12 yr after strobe rearing. The deficits were remarkably specific; strobe rearing did not affect spatial receptive-field structure, orientation selectivity, spatial or temporal frequency tuning, or general responsiveness to visual stimuli. These results provide further support for a critical role of S-T structure in determining direction selectivity in simple cells. Strobe rearing eliminates directional tuning by altering the timing of responses within the receptive field.     

Spike train encoding by regular-spiking cells of the visual cortex

1. To study the encoding of input currents into output spike trains by regular-spiking cells, we recorded intracellularly from slices of the guinea pig visual cortex while injecting step, sinusoidal, and broadband noise currents. 2. When measured with sinusoidal currents, the frequency tuning of the spike responses was markedly band-pass. The preferred frequency was between 8 and 30 Hz, and grew with stimulus amplitude and mean intensity. 3. Stimulation with broadband noise currents dramatically enhanced the gain of the spike responses at low and high frequencies, yielding an essentially flat frequency tuning between 0.1 and 130 Hz. 4. The averaged spike responses to sinusoidal currents exhibited two nonlinearities: rectification and spike synchronization. By contrast, no nonlinearity was evident in the averaged responses to broadband noise stimuli. 5. These properties of the spike responses were not present in the membrane potential responses. The latter were roughly linear, and their frequency tuning was low-pass and well fit by a single-compartment passive model of the cell membrane composed of a resistance and a capacitance in parallel (RC circuit). 6. To account for the spike responses, we used a "sandwich model" consisting of a low-pass linear filter (the RC circuit), a rectification nonlinearity, and a high-pass linear filter. The model is described by six parameters and predicts analog firing rates rather than discrete spikes. It provided satisfactory fits to the firing rate responses to steps, sinusoids, and broadband noise currents. 7. The properties of spike encoding are consistent with temporal nonlinearities of the visual responses in V1, such as the dependence of response frequency tuning and latency on stimulus contrast and bandwidth. We speculate that one of the roles of the high-frequency membrane potential fluctuations observed in vivo could be to amplify and linearize the responses to lower, stimulus-related frequencies.     

Synchronization of visual responses between the cortex, lateral geniculate nucleus, and retina in the anesthetized cat

Synchronization of spatially distributed responses in the cortex is often associated with periodic activity. Recently, synchronous oscillatory patterning was described for visual responses in retinal ganglion cells that is reliably transmitted by the lateral geniculate nucleus (LGN), raising the question of whether oscillatory inputs contribute to synchronous oscillatory responses in the cortex. We have made simultaneous multi-unit recordings from visual areas 17 and 18 as well as the LGN and the retina to examine the interactions between subcortical and cortical synchronization mechanisms. Strong correlations of oscillatory responses were observed between retina, LGN, and cortex, indicating that cortical neurons can become synchronized by oscillatory activity relayed through the LGN. This feedforward synchronization occurred with oscillation frequencies in the range of 60-120 Hz and was most pronounced for responses to stationary flashed stimuli and more frequent for cells in area 18 than in area 17. In response to moving stimuli, by contrast, subcortical and cortical oscillations dissociated, proving the existence of independent subcortical and cortical mechanisms. Subcortical oscillations maintained their high frequencies but became transient. Cortical oscillations were now dominated by a cortical synchronizing mechanism operating in the 30-60 Hz frequency range. When the cortical mechanism dominated, LGN responses could become phase-locked to the cortical oscillations via corticothalamic feedback. In summary, synchronization of cortical responses can result from two independent but interacting mechanisms. First, a transient feedforward synchronization to high-frequency retinal oscillations, and second, an intracortical mechanism, which operates in a lower frequency range and induces more sustained synchronization.     

Effects of saccades on the activity of neurons in the cat lateral geniculate nucleus

Effects of saccades on individual neurons in the cat lateral geniculate nucleus (LGN) were examined under two conditions: during spontaneous saccades in the dark and during stimulation by large, uniform flashes delivered at various times during and after rewarded saccades made to small visual targets. In the dark condition, a suppression of activity began 200-300 ms before saccade start, peaked approximately 100 ms before saccade start, and smoothly reversed to a facilitation of activity by saccade end. The facilitation peaked 70-130 ms after saccade end and decayed during the next several hundred milliseconds. The latency of the facilitation was related inversely to saccade velocity, reaching a minimum for saccades with peak velocity >70-80 degrees /s. Effects of saccades on visually evoked activity were remarkably similar: a facilitation began at saccade end and peaked 50-100 ms later. When matched for saccade velocity, the time courses and magnitudes of postsaccadic facilitation for activity in the dark and during visual stimulation were identical. The presaccadic suppression observed in the dark condition was similar for X and Y cells, whereas the postsaccadic facilitation was substantially stronger for X cells, both in the dark and for visually evoked responses. This saccade-related regulation of geniculate transmission appears to be independent of the conditions under which the saccade is evoked or the state of retinal input to the LGN. The change in activity from presaccadic suppression to postsaccadic facilitation amounted to an increase in gain of geniculate transmission of approximately 30%. This may promote rapid central registration of visual inputs by increasing the temporal contrast between activity evoked by an image near the end of a fixation and that evoked by the image immediately after a saccade.     

Representation of spectral and temporal sound features in three cortical fields of the cat. Similarities outweigh differences

This study investigates the degree of similarity of three different auditory cortical areas with respect to the coding of periodic stimuli. Simultaneous single- and multiunit recordings in response to periodic stimuli were made from primary auditory cortex (AI), anterior auditory field (AAF), and secondary auditory cortex (AII) in the cat to addresses the following questions: is there, within each cortical area, a difference in the temporal coding of periodic click trains, amplitude-modulated (AM) noise bursts, and AM tone bursts? Is there a difference in this coding between the three cortical fields? Is the coding based on the temporal modulation transfer function (tMTF) and on the all-order interspike-interval (ISI) histogram the same? Is the perceptual distinction between rhythm and roughness for AM stimuli related to a temporal versus spatial representation of AM frequency in auditory cortex? Are interarea differences in temporal response properties related to differences in frequency tuning? The results showed that: 1) AM stimuli produce much higher best modulation frequencies (BMFs) and limiting rates than periodic click trains. 2) For periodic click trains and AM noise, the BMFs and limiting rates were not significantly different for the three areas. However, for AM tones the BMF and limiting rates were about a factor 2 lower in AAF compared with the other areas. 3) The representation of stimulus periodicity in ISIs resulted in significantly lower mean BMFs and limiting rates compared with those estimated from the tMTFs. The difference was relatively small for periodic click trains but quite large for both AM stimuli, especially in AI and AII. 4) Modulation frequencies <20 Hz were represented in the ISIs, suggesting that rhythm is coded in auditory cortex in temporal fashion. 5) In general only a modest interdependence of spectral- and temporal-response properties in AI and AII was found. The BMFs were correlated positively with characteristic frequency in AAF. The limiting rate was positively correlated with the frequency-tuning curve bandwidth in AI and AII but not in AAF. Only in AAF was a correlation between BMF and minimum latency was found. Thus whereas differences were found in the frequency-tuning curve bandwidth and minimum response latencies among the three areas, the coding of periodic stimuli in these areas was fairly similar with the exception of the very poor representation of AM tones in AII. This suggests a strong parallel processing organization in auditory cortex.     

An attempt to give nursing home residents a voice in the quality improvement process: the challenge of frailty

A model for the development of spatiotemporal receptive fields of simple cells in the visual cortex is proposed. The model is based on the 1990 hypothesis of Saul and Humphrey that the convergence of four types of input onto a cortical cell, viz. non-lagged ON and OFF inputs and lagged ON and OFF inputs, underlies the spatial and temporal structure of the receptive fields. It therefore explains both orientation and direction selectivity of simple cells. The response properties of the four types of input are described by the product of linear spatial and temporal response functions. Extending the 1994 model of one of the authors (K.D. Miller), we describe the development of spatiotemporal receptive fields as a Hebbian learning process taking into account not only spatial but also temporal correlations between the different inputs. We derive the correlation functions that drive the development both for the period before and after eye-opening and demonstrate how the joint development of orientation and direction selectivity can be understood in the framework of correlation-based learning. Our investigation is split into two parts that are presented in two papers. In the first, the model for the response properties and for the development of direction-selective receptive fields is presented. In the second paper we present simulation results that are compared with experimental data, and also provide a first analysis of our model.     

Functional organization of owl monkey lateral geniculate nucleus and visual cortex

The nocturnal, New World owl monkey (Aotus trivirgatus) has a rod-dominated retina containing only a single cone type, supporting only the most rudimentary color vision. However, it does have well-developed magnocellular (M) and parvocellular (P) retinostriate pathways and striate cortical architecture [as defined by the pattern of staining for the activity-dependent marker cytochrome oxidase (CO)] similar to that seen in diurnal primates. We recorded from single neurons in anesthetized, paralyzed owl monkeys using drifting, luminance-modulated sinusoidal gratings, comparing receptive field properties of M and P neurons in the lateral geniculate nucleus and in V1 neurons assigned to CO "blob," "edge," and "interblob" regions and across layers. Tested with achromatic stimuli, the receptive field properties of M and P neurons resembled those reported for other primates. The contrast sensitivity of P cells in the owl monkey was similar to that of P cells in the macaque, but the contrast sensitivities of M cells in the owl monkey were markedly lower than those in the macaque. We found no differences in eye dominance, orientation, or spatial frequency tuning, temporal frequency tuning, or contrast response for V1 neurons assigned to different CO compartments; we did find fewer direction-selective cells in blobs than in other compartments. We noticed laminar differences in some receptive field properties. Cells in the supragranular layers preferred higher spatial and lower temporal frequencies and had lower contrast sensitivity than did cells in the granular and infragranular layers. Our data suggest that the receptive field properties across functional compartments in V1 are quite homogeneous, inconsistent with the notion that CO blobs anatomically segregate signals from different functional "streams."     

Coding of the acoustic information in the superior auditory centers

The internal ear may be considered analysing acoustical signals in the frequency domain. This spectral analysis appears in the auditory pathways as a place code, each neuron being activated for a narrow and well defined frequency band. But in addition to this place code, temporal information on the phase and the period of low frequency signals is preserved in the low auditory centers. In the medial geniculate body, the last relay before the cerebral cortex, the place code shows the same properties as in lower centers but with a greater diversity in the response patterns and tuning properties. The tonotopic organization is less precise and, for the pars lateralis, follows the histological lamellar organization of this region. The most lateral laminae are composed of cells responding to low frequencies, the most medial ones of high frequency cells. In the auditory cortex intracellular recordings confirm the importance of an active inhibition underlying the diverse response patterns observed. Persistance of a time code is shown by certain cells presenting responses precisely time-locked to individual clicks in a train for rates ranging from 50 to 1000 Hz. Other cells respond selectively for certain click train frequencies without marking the temporal structure of the stimuli. Thus a temporal and a place code are still both present at the cortical level for this particular kind of signals.     

Spatiotemporal receptive field organization in the lateral geniculate nucleus of cats and kittens

We have studied the spatiotemporal receptive-field organization of 144 neurons recorded from the dorsal lateral geniculate nucleus (dLGN) of adult cats and kittens at 4 and 8 wk postnatal. Receptive-field profiles were obtained with the use of a reverse correlation technique, in which we compute the cross-correlation between the action potential train of a neuron and a randomized sequence of long bright and dark bar stimuli that are flashed throughout the receptive field. Spatiotemporal receptive-field profiles of LGN neurons generally exhibit a biphasic temporal response, as well as the classical center-surround spatial organization. For nonlagged cells, the first temporal phase of the response dominates, whereas for lagged neurons, the second temporal phase of the response is typically the largest. This temporal phase difference between lagged and nonlagged cells accounts for their divergent behavior in response to flashed stimuli. Most LGN cells exhibit some degree of space-time inseparability, which means that the receptive field cannot simply be viewed as the product of a spatial waveform and a temporal waveform. In these cases, the response of the surround is typically delayed relative to that of the center, and there is some blending of center and surround during the time course of the response. We demonstrate that a simple extension of the traditional difference-of-Gaussians (DOG) model, in which the surround response is delayed relative to that of the center, accounts nicely for these findings. With regard to development, our analysis shows that spatial and temporal aspects of receptive field structure mature with markedly different time courses. After 4 wk postnatal, there is little change in the spatial organization of LGN receptive fields, with the exception of a weak, but significant, trend for the surround to become smaller and stronger with age. In contrast, there are substantial changes in temporal receptive-field structure after 4 wk postnatal. From 4 to 8 wk postnatal, the shape of the temporal response profile changes, becoming more biphasic, but the latency and duration of the response remain unchanged. From 8 wk postnatal to adulthood, the shape of the temporal profile remains approximately constant, but there is a dramatic decline in both the latency and duration of the response. Comparison of our results with recent data from cortical (area 17) simple cells reveals that the temporal development of LGN cells accounts for a substantial portion of the temporal maturation of simple cells.     

Temporal aspects of the effects of cooling on responses of single auditory nerve fibers

During an investigation of the effects of cochlear cooling on frequency tuning and input/output relations of single auditory nerve fibers in gerbil (Ohlemiller and Siegel (1994) Hear. Res. 80, 174-190), cooling-related changes in post-stimulus time histogram (PSTH) shape and phase-locking to tonebursts were characterized in a small sample of neurons. Local cochlear cooling by 5-10 degrees C below normal core temperature did not alter overall PSTH shape, although some evidence was found for a reduction in the time constants of rapid and short term rate adaptation. The relative contributions of rapid and short term response components appeared unaltered. Effects of cooling on phase-locking were assessed by calculating the synchronization index for responses to intense ( > 70 dB SPL) tonebursts at 0.5, 1.0, and 2.0 kHz. Synchronization filter functions exhibited modest reductions in both magnitude and the upper frequency limit of phase-locking. The effects of cooling on the temporal character of responses appear distinct from those of a simple reduction in stimulus intensity. Results are interpreted in terms of cooling-related changes in responses of cochlear hair cells and afferent neurons, and suggest that temperature artifacts are unlikely to underlie reported species differences in PSTH shape and phase-locking.     

The intrinsic temporal properties of alpha and beta retinal ganglion cells are equivalent

BACKGROUND: Mammalian retinal ganglion cells have been traditionally classified on the basis of morphological and functional criteria, but as yet little is known about the intrinsic membrane properties of these neurons. This study has investigated these properties by making patch-clamp recordings from morphologically identified ganglion cells in the intact retina. RESULTS: The whole-cell configuration of the patch-clamp technique was used to assess the temporal tuning characteristics of alpha and beta cells, the two most extensively studied ganglion cell classes. Fourier analysis was used to examine discharge patterns in response to sinusoidal currents of different frequencies (1-50 Hz). With few exceptions, neurons responded in a stereotypic fashion to changes in temporal modulation, with their output initially increasing and then decreasing as a function of stimulus frequency. Moreover, peak responses in both cell classes were obtained at equivalent temporal frequencies. At high stimulus rates, response probability decreased, but the spikes remained phase-locked to the stimulus cycle, thereby enabling populations of cells to convey temporal information. A small number of ganglion cells did not show an appreciable decrease in output as a function of stimulus frequency, but these cells were not confined to either ganglion cell class. CONCLUSIONS: These findings provide the first evidence that the intrinsic temporal properties of alpha and beta cells are alike. Furthermore, the responses obtained to direct current injections were strikingly similar to those described previously with temporally modulated visual stimuli, suggesting that intrinsic membrane properties may shape the visual responses of alpha and beta cells to a larger degree than has been commonly assumed.     

Acoustic distortion as a measure of frequency selectivity: relation to psychophysical equivalent rectangular bandwidth

The magnitude of cubic intermodulation distortion generated when two tones are progressively separated in frequency reaches a broad maximum when the distortion frequency falls just over half an octave below the high-frequency stimulus (f2), when this distortion is measured with a microphone in the ear canal. For the component 2f1-f2, this peak occurs at an f2/f1 ratio of approximately 1.2. The tuning, magnitude, and mean group delay of this distortion peak was measured for a fixed f2 of 4 kHz at 40 dB SPL and a varied f1 at 55 dB SPL in eight human subjects with normal hearing. The distortion peak measures were compared with the frequency selectivity at 4 kHz of the same eight subjects derived using a forward-masking notched-noise paradigm. In the six subjects from whom good, repeatable levels of distortion were measured, a significant negative correlation was found between the tuning of the distortion peak and the psychophysical bandwidth at f2. It is concluded that the tuning of the distortion peak may provide an objective measure of frequency selectivity in the human cochlea.     

Processing of first- and second-order motion signals by neurons in area MT of the macaque monkey

Extrastriate cortical area MT is thought to process behaviorally important visual motion signals. Psychophysical studies suggest that visual motion signals may be analyzed by multiple mechanisms, a "first-order" one based on luminance, and a "second-order" one based upon higher level cues (e.g. contrast, flicker). Second-order motion is visible to human observers, but should be invisible to first-order motion sensors. To learn if area MT is involved in the analysis of second-order motion, we measured responses to first- and second-order gratings of single neurons in area MT (and in one experiment, in area V1) in anesthetized, paralyzed macaque monkeys. For each neuron, we measured directional and spatio-temporal tuning with conventional first-order gratings and with second-order gratings created by spatial modulation of the flicker rate of a random texture. A minority of MT and V1 neurons exhibited significant selectivity for direction or orientation of second-order gratings. In nearly all cells, response to second-order motion was weaker than response to first-order motion. MT cells with significant selectivity for second-order motion tended to be more responsive and more sensitive to luminance contrast, but were in other respects similar to the remaining MT neurons; they did not appear to represent a distinct subpopulation. For those cells selective for second-order motion, we found a correlation between the preferred directions of first- and second-order motion, and weak correlations in preferred spatial frequency. These cells preferred lower temporal frequencies for second-order motion than for first-order motion. A small proportion of MT cells seemed to remain selective and responsive for second-order motion. None of our small sample of V1 cells did. Cells in this small population, but not others, may perform "form-cue invariant" motion processing (Albright, 1992).     

The effect of noise exposure on the details of distortion product otoacoustic emissions in humans

The effect of noise exposure on amplitude and phase of distortion product otoacoustic emissions (DPOAEs) was examined by five different paradigms: across a wideband of frequency, microstructure, input/output function, primary frequency ratio tuning curve, and group delay. The aim was to investigate the vulnerability of these different features to moderate levels of noise exposure. Nine subjects were exposed to third-octave-band noise. The DPOAE amplitude was reduced frequency specifically with the greatest reduction approximately half an octave above the frequency of the noise. The degree of amplitude reduction was greatest at low stimulus levels. There were no observed effects on the shape of the primary ratio tuning curve. A weak tendency to a decrease was seen in group delays. Distinct microstructure was seen in the amplitude against frequency of five out of seven subjects. The maximum to minimum ratio of the microstructure decreased, and the whole pattern shifted toward lower frequencies after noise exposure. Evidence of multiple internal reflection or interference was seen in the periodicity of the microstructure. Using a simple model of the microstructure based on multiple reflections, the noise-induced changes were reevaluated. A reduction in maximum to minimum microstructure ratio could be interpreted as a decrease in the internal reflection coefficient. The implications of these observations for the interpretation of the DPOAE measurements are considered.     

The effect of age on the reversibility of cellular atrophy in the LGN of the cat following monocular deprivation: a test of two hypotheses about cell growth

Monocular closure soon after birth is known to reduce the growth of the deprived cells in the LGN provided that there are competing cells with normal input. When the closed eye was opened and the open eye closed after three of six weeks of monocular closure, the originally deprived cells in the LGN were able to recover their normal size. However, it was found that after 14 weeks of monocular closure there was no recovery. The ability of the initially closed eye to excite cells in the visual cortex is known to depend on the age when eye closure is reversed in a similar manner. Thus the growth of cells in the LGN is correlated with the possession of effective synapses upon cortical cells. This result is compatible with the possession of effective synapses upon cortical cells. This result is compatible with the hypothesis that competition occurs at the cortical level. It is argued that the alternative hypothesis of competition within the LGN predicts reversibility at any age. In some kittens, the closed eye was opened and the opposite optic nerve crushed. Some evidence was then found of structural recovery even after 14 weeks of deprivation. The hypothesis of cortical competition predicts functional recovery in such kittens, but this remains to be tested.     

Plasticity of neuronal response properties in adult cat striate cortex

We have utilized an associative conditioning paradigm to induce changes in the receptive field (RF) properties of neurons in the adult cat striate cortex. During conditioning, the presentation of particular visual stimuli were repeatedly paired with the iontophoretic application of either GABA or glutamate to control postsynaptic firing rates. Similar paradigms have been used in kitten visual cortex to alter RF properties (Fregnac et al., 1988, 1992; Greuel et al., 1988; Shulz & Fregnac, 1992). Roughly half of the cells that were subjected to conditioning with stimuli differing in orientation were found to have orientation tuning curves that were significantly altered. In general, the modification in orientation tuning was not accompanied by a shift in preferred orientation, but rather, responsiveness to stimuli at or near the positively reinforced orientation was increased relative to controls, and responsiveness to stimuli at or near the negatively reinforced orientation was decreased relative to controls. A similar proportion of cells that were subjected to conditioning with stimuli differing in spatial phase were found to have spatial-phase tuning curves that were significantly modified. Conditioning stimuli typically differed by 90 deg in spatial phase, but modifications in spatial-phase angle were generally 30-40 deg. An interesting phenomenon we encountered was that during conditioning, cells often developed a modulated response to counterphased grating stimuli presented at the null spatial phase. We present an example of a simple cell for which the shift in preferred spatial phase measured with counterphased grating stimuli was comparable to the shift in spatial phase computed from a one-dimensional Gabor fit of the space-time RF profile. One of ten cells tested had a significant change in direction selectivity following associative conditioning. The specific and predictable modifications of RF properties induced by our associative conditioning procedure demonstrate the ability of mature visual cortical neurons to alter their integrative properties. Our results lend further support to models of synaptic plasticity where temporal correlations between presynaptic and postsynaptic activity levels control the efficiency of transmission at existing synapses, and to the idea that the mature visual cortex is, in some sense, dynamically organized.