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Group in Vision Science, School of Optometry, and Helen Wills Neuroscience Institute, University of California, Berkeley, California 94720-2020
Submitted 13 June 2003; accepted in final form 29 October 2003
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMETHODSRESULTSDISCUSSIONACKNOWLEDGMENTSREFERENCES |
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMETHODSRESULTSDISCUSSIONACKNOWLEDGMENTSREFERENCES |
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; Ohzawa et al. 1990; Qian and Andersen 1994; Qian et al. 1994).
An early theoretical proposal was concerned with the correspondence problem in stereopsis (Marr and Poggio 1979). This refers to the ambiguity of correspondence between left and right images that occurs from binocular viewing. The brain must choose the correct depth plane from several possible ones to process appropriate stereoscopic information. To solve this problem, coarse scale disparity matches were proposed to occur first. This would be followed by fine scale matches (Marr and Poggio 1979). In other words, a coarse-to-fine scaling process could be used to provide correct stereoscopic matching. Other theoretical ideas envisioned similar coarse scale disparity matches that were followed by fine scale adjustments (Anderson and Van Essen 1987; Nishihara and Kimura 1987; Quam 1987).
These theoretical notions have been explored in behavioral studies. Sensitivity has been found to improve for line length, orientation, curvature and stereoscopic depth over a period of 
1 s (Watt 1987). The interpretation of these findings may be made in terms of a coarse-to-fine temporal analysis of spatial features. Another study was aimed at determining if different spatial scales interact in stereopsis (Rohaly and Wilson 1993; Wilson et al. 1991). Diplopia threshold was determined at two separate spatial scales. Results suggested that coarse scale disparity processing constrains that of fine levels within a given range. Two experiments were performed in another study in which spatially filtered targets were used. In one, temporal sequences were shown in which full-bandwidth targets were compared with those in which selected frequencies were presented first. In a second experiment, a human face was used in a similar way. Results of both experiments provide clear evidence for anisotropic temporal processing. Specifically, the most efficient perceptual processing occurs when spatial information is presented temporally in a low-to-high spatial frequency sequence. (Parker et al. 1997)
Other psychophysical investigations of stereopsis suggest that there is also a fine-to-coarse process. In one study, an ambiguous coarse scale stimulus was presented that could be perceived with either crossed or uncrossed disparity. When a fine scale stimulus was added, the ambiguity at coarse scale was removed, suggesting a fine-to-coarse disambiguation process (Smallman 1995). However, this study also showed that a coarse scale stimulus could disambiguate that of a fine scale. Therefore results of the study support both coarse-to-fine and fine-to-coarse processes.
Considered together, results of the behavioral studies suggest that both coarse-to-fine and fine-to-coarse processes may apply to stereopsis. Surprisingly, until our recent study (Menz and Freeman 2003), no physiological data on this issue have been available. Neuronal temporal analysis on a fine time scale has become possible with relatively recent techniques such as reverse correlation analysis (DeAngelis et al. 1993a,b; Freeman and Ohzawa 1990; Jones and Palmer 1987). Results of orientation (Ringach et al. 1997) and spatial frequency (Bredfeldt and Ringach 2002) tuning studies using this technique show prominent changes as responses evolve over time. This type of information is important because it provides clues about neural circuitry. In ideal cases, for example, it may be possible to obtain evidence consistent with feed-forward or feedback models of visual processing.
We have carried out a temporal analysis over a brief time scale (i.e., 40 ms) for a population of neurons in lateral geniculate nucleus (LGN) and visual cortex. In addition to determining specific temporal features relevant to stereoscopic processing, our aim was to obtain information concerning the theoretical proposal of a coarse-to-fine sequence as outlined in the preceding text. Our results provide clear evidence that is consistent with this hypothesis.
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METHODS |
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TOPABSTRACTINTRODUCTIONMETHODSRESULTSDISCUSSIONACKNOWLEDGMENTSREFERENCES |
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,b). The description provided here emphasizes specific procedures that are relevant to this study. Surgical procedures and animal maintenance
Following standard preanesthetic procedures, isoflurane is used to anesthetize the animal. Femoral veins are cannulated, a tracheal tube is placed, and a craniotomy and durotomy are performed (H-C P4 L2 for visual cortex and A6 L8 for LGN). After surgery, thiopental is used to maintain anesthesia. Each animal is assessed individually to determine an adequate level of anesthesia, which generally ranges from 
1.5–2.5 mg · kg–1 · h–1. After anesthesia level is stabilized over 1 h, a muscle relaxant (gallamine triethiodide, 10 mg · kg–1 · h–1) is used to prevent eye movements during visual stimulation. Pupils are dilated, nictitating membranes are retracted, and contact lenses with 4-mm artificial pupils are positioned. A reversible direct ophthalmoscope is used to image the optic discs on a tangent screen to infer areae centrales locations (Bishop et al. 1962). Core body temperature, electroencephalogram (EEG), electrocardiogram (ECG), heart rate, intratracheal pressure, and expired CO2 are all monitored continuously throughout each experiment.
Recording procedure
Visual stimuli are generated by a computer with two high-resolution graphics boards that runs custom software as described previously (DeAngelis et al. 1993a,b). To map cortical RFs, a dichoptic one-dimensional binary m-sequence noise stimulus is used (Anzai et al. 1999a,b). A monocular stimulus is used for LGN cells. Sixteen long adjacent bars are presented to each eye at optimal orientation (Fig. 1A). The width of the bars is approximately one-fourth the period of the optimal frequency. This square pattern is centered over the RF. Each of the 16 bars is either bright or dark, and the background is the mean luminance of the bars.
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) to obtain space-time RF maps (Fig. 1B). This is repeated for all time delays of interest (0–200 ms) in increments of 5 ms to obtain a space-time RF. A nonlinear binocular interaction map is obtained by cross-correlating the spike train with each spatial location in the binocular view field pattern. In the maps shown in Fig. 1, C and D, the white and dark regions indicate excitatory responses to same or opposite polarity bars, respectively. Data analysis
Our analysis smoothes or interpolates the RFs within the 40-ms duration stimuli and uses eight bins to achieve space-time RFs with 5-ms interval correlation delays. It is possible that the autocorrelation of the stimulus produces an artifactual dynamic (Theunissen et al. 2001). To test for this possibility, we also generated RFs using bins equal in size to the stimulus duration (40 ms). The two-dimensional binocular interaction maps are reduced to one-dimensional disparity-tuning data by integrating along lines of equal disparity (Ohzawa et al. 1997) (see Fig. 1, C and D). These data are fit with a Gabor function by the Levenberg-Marquardt algorithm (Press et al. 1992)
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s is the width or size parameter, fds is the disparity frequency, and 
is the phase. For LGN cells, the fit in the spatial domain is a single Gaussian function
 | (2) |
c is the size parameter. A difference of Gaussians (DOG) fit was attempted, but the surround was so weak at nonoptimal time slices that this procedure was not practical. For the purpose of this study, a single Gaussian function is suitable. The most direct, assumption-free method of analyzing frequency content is to take the Fourier Transform at each time delay and examine the change in optimal frequency and bandwidth. The frequency data were fit with a Gaussian function and the dynamics measured this way closely match the method of fitting a Gabor function in the spatial domain (data not shown).
At each time slice, the value of a parameter is normalized (i.e., divided) by the value at the optimal time slice. A linear regression is performed on the parameters as a function of time delay relative to optimal. The slope is used as a measure of the rate of change of the parameter. The value from the regression is multiplied by 1,000 yielding units of %/10 ms. For the monocular RFs of disparity-tuned simple cells, a separate regression is performed for the left and right eye findings, and the two numbers are averaged to obtain an overall monocular result. Unless otherwise noted, statistics are based on a standard normal distribution as described by the Central Limit theorem, which requires a large sample size.
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RESULTS |
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TOPABSTRACTINTRODUCTIONMETHODSRESULTSDISCUSSIONACKNOWLEDGMENTSREFERENCES |
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; Skottun et al. 1991). Although LGN cells are monocular, except for modest dichoptic effects, we include them in this study to gain insights into possible mechanisms of organization of the neural system for disparity processing. Our main focus is an examination of temporal response characteristics of each cell type in the visual cortex. We examine both monocular and binocular properties of simple cells to compare temporal properties in the processing sequence. For LGN cells, we examine changes in RF size during responses. We use best-fitting Gaussian functions to make this assessment. For cortical cells, best-fitting Gabor functions are used and the variable of interest is correlation delay. LGN
We first consider the spatial properties and temporal dynamics of LGN cells. We assume a serial processing system in which input is fed from LGN neurons to simple and then to complex cells. The actual nature of this system is not crucial for our analysis, and we assume that there is also parallel processing.
An example of temporal characteristics of the RF profile for an LGN cell is shown in Fig. 2. The data in A–C are fit with single Gaussian functions. They show, respectively, RF profiles for 15 ms before the optimal delay, at the optimal value, and 15 ms after optimal delay. Widths at half-maximum amplitudes are designated (
). Normalized RF width as a function of time delay, given in Fig. 2D, shows a clear reduction in RF size with increasing correlation delay. For this cell, the center size decreases at a rate of 15.4%/10 ms. Note that the range of usable time delays is relatively small because the duration of the first phase of the temporal response is short. This is due to LGN preference for high temporal frequencies. Note also that in the example shown in Fig. 2, LGN center-surround organization is not evident in the early part of the response (A) but it appears weakly at a subsequent time slice (C). This result is consistent with our previous work showing that LGN surrounds are frequently time-delayed relative to the center response (Cai et al. 1997).
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). For a time-delayed surround, the center RF subregion decreases in size with correlation delay. Overall RF size, which includes the surround, increases with correlation delay. The combination of RF center size decrease with overall size increase can account for increased frequency (resolution) and increased overall size of monocular RFs of simple cells.
Population data for the LGN cell recordings are given in Fig. 3. The histograms here show the change in size of the RF center during the temporal windows indicated. The distribution has an approximately Gaussian shape, but it includes a subgroup of cells (from –20 to –16%/10 ms) that exhibits a relatively large rate of decrease in RF center size. The average rate of decrease for the entire sample is 7.6%/10 ms. This is significantly different from no change (P < 0.0001). It is a relatively large effect compared with that for cells in the visual cortex and is greater by 3–4%/10 ms (see following text).
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An example of disparity-tuning dynamics of a simple cell is shown in Fig. 5. Binocular components are given in Fig. 5, A–D. Monocular functions are depicted in E–L. Each set of data points for temporal slices at optimal and ±25 ms before and after optimal, are fitted with Gabor functions (Fig. 5, A–C). Inspection of these functions shows that flanking subregions move closer to the central peak as correlation delay increases. The position of the central peak remains constant for all delay times. At higher correlation delays, the RF subregions decrease in size and move closer together. These changes are expressed in terms of the Gabor function fits in disparity frequency (resolution) and size (range) in Fig. 5D. Data points, presented for 5-ms differences from optimal time (see following text), show a linear decrease and increase for size and frequency, respectively.
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Complex cells
The disparity-tuning dynamics of a complex cell are illustrated in Fig. 6. As in the previous example, tuning curves, fit with Gabor functions, are shown for temporal delays at the optimal time (B) and at –20 and +20 ms (A and C, respectively). Vertical lines mark the peak and troughs at the +20 ms level (C). The vertical bars are extended into A and B to help show that the flanking RF subregions move closer to the central peak with increasing correlation delay. Subregions also become smaller and closer together. Note that for all 3 time delays, the position of the central peak remains constant. From the Gabor function fits, RF disparity frequency (resolution) and disparity range (size) may be estimated. The data (Fig. 6D) show that RF range (size) decreases approximately linearly with increasing time delay. A similar but opposite change is shown for frequency (resolution) that increases with increasing time delay. The results of Fig. 6 show clearly that optimal disparity, defined by the location of the main peak, does not change with correlation delay. This finding, combined with the increase in disparity frequency (resolution) and decrease in disparity range (size) constitutes a coarse-to-fine disparity process. This kind of mechanism was put forward in a model that proposed to account for how the visual system solves the retinal disparity correspondence problem (Marr and Poggio 1979). In the following text, we examine our cell population to see if the overall results are consistent with this notion.
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). This method produces similar results as we show in the following text. In a previous report, these issues have been clearly discussed (Ringach et al. 2003)
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and 
, respectively). Distributions for the two binning methods are broadly similar and not significantly different (P = 0.93 and P = 0.42 Wilcoxon rank-sum test, for frequency and range distributions, respectively). This finding confirms the justification to use the shorter interpolated bin width, and subsequent data are presented in this form because it provides a larger sample size. For the entire distribution using the 5-ms binning method (Fig. 8A), average increase in disparity frequency is 4.5%/10 ms. This is a relatively modest but clear effect. Only 9 of 87 cells show decreases in frequency, and they are relatively small. The data for disparity range change, presented in Fig. 8B, also shows a clear trend, but it is weaker than that for frequency. The average decrease in disparity range is 3.2%/10 ms. In the case of range, 17 of 87 cells show a reverse effect, i.e., an increase rather than a decrease.
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A primary question of this study, posed at the outset, is related to the notion of coarse-to-fine processing. One relevant parameter is the relation between optimal time delay and different stimulus parameters. Specifically, in a coarse-to-fine process, optimal time delays could be longer for high compared with low disparity frequencies. This means that coarse information would be processed first followed by that of fine detail. A comparison of optimal time delay and disparity frequency for our population of simple and complex cells is shown in Fig. 9, A and B, respectively. Although the distributions are relatively broad, there is a clear tendency, for both simple and complex cells, for optimal time delay to increase with disparity frequency. Robust regression lines fit to the data have slopes of 0.0081 and 0.0085 cycle · °–1 · ms–1 for simple (A) and complex (B) cells, respectively. These values are significantly different from zero (P = 0.031 and 0.0001, respectively). Clearly, neurons with higher preferred disparity frequencies are relatively more time delayed. The correlation is slightly stronger for complex compared with simple cells (correlation coefficients of 0.42 and 0.33, respectively). This coarse-to-fine dynamics could be accounted for by neurons that pool input from other cells of slightly different disparity frequency content. The lower disparity frequency information is represented relatively earlier in the response. Our data for both complex and simple cells are consistent with this feed-forward mechanism for generating coarse-to-fine disparity tuning.
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DISCUSSION |
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TOPABSTRACTINTRODUCTIONMETHODSRESULTSDISCUSSIONACKNOWLEDGMENTSREFERENCES |
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). Note, however, that the coarse-to-fine mechanism that we have investigated covers a relatively narrow range. To eliminate false matches, this process requires a considerable narrowing of the range. It is possible that other mechanisms beyond V1 effectively expand the range of this function. In any case, our results leave open the possibility that this coarse-to-fine process is not restricted to the correspondence problem. It may apply to the entire stereoscopic process, and it also could underlie other visual functions. Visual perceptions could involve temporal sequences by which low spatial frequency information is processed first providing an initial coarse view which is then refined with time. The mechanism for this process could begin at or prior to the LGN. Our present findings show that LGN cells exhibit a decrease in center size with correlation delay. This result is associated with a time-delayed LGN RF surround. This effect could contribute to the increased Gabor function spatial frequency with time that is exhibited in monocular RFs of simple cells. In general, disparity dynamics of simple and complex cells are similar. This could be the result of pooled input from simple to complex cells. However, there is a discrepancy between monocular and binocular dynamics of disparity-sensitive simple cells. Disparity frequency dynamics are similar in the two conditions but range (size) temporal changes occur in opposite directions, i.e., monocular RFs increase in size while disparity range (size) decreases. This effect is considered below.
Tuning dynamics
Important theoretical concepts have been put forward to account for motion detection and related perceptual phenomena. Energy models may be most suitable for this purpose (Adelson and Bergen 1985; Watson and Ahumada 1985). These models have been modified to provide a theoretical basis for the processing of stereoscopic information (Anzai et al. 2001; DeAngelis et al. 1995; Freeman and Ohzawa 1990; Ohzawa et al. 1997), and experimental tests have been conducted to determine if predictions of the models are matched by the resulting data. Most of these tests are concerned with changes in spatial location of a subregion over time. In the work reported here, we are concerned with changes in tuning characteristics over a relatively limited time epoch (40 ms).
Previous studies have been conducted in which the dynamics of tuning characteristics of cortical cells have been examined. The most common characteristic is orientation tuning. Results of these studies are mixed. Orientation bandwidth has been reported to become narrower during the temporal course of the response of cortical cells (Best et al. 1989; Ringach et al. 1997; Volgushev et al. 1995). In other experimental work, orientation tuning characteristics are reported to be largely unchanged during the temporal course of the response (Celebrini et al. 1993; Mazer et al. 2002). The spatially changing positive effects were from anesthetized preparations and the time-constant findings were from awake behaving animals. It is not clear if this difference is relevant to the findings. The question of temporal dynamics has also been applied to spatial frequency tuning. In this case, it has been reported that tuning preference changes from lower to higher spatial frequencies over time (Bredfeldt and Ringach 2000). This latter finding is consistent with our current results. Finally, a recently reported study of spatial dynamics of RFs in visual cortex utilized relatively long-duration stimuli (300 ms). The reported finding is that RF subregion width decreased with greater delays in a reverse correlation procedure (Suder et al. 2002). Their data are more consistent with a feed–forward model from thalamic input than with one that includes intracortical feedback. A feed–forward mechanism is what we propose to account for the results of the current study.
In addition to the type of preparation question noted in the preceding text, there are other experimental differences in the approach to this problem. For the recent studies, a type of reverse correlation stimulus procedure has been used (DeAngelis et al. 1993a,b). Generally, this means that a spike train is cross-correlated to a noise stimulus sequence. In the case of our present study, the analysis is based on a temporal bin size that is equal to the stimulus duration. Our results of coarse-to-fine tuning are clear and consistent across the population we have studied. The other relevant result of our study is that monocular and binocular RFs of simple cells change size in opposite directions with time. This would not occur if the basic result was due to an artifactual dynamic.
Mechanisms
The original qualitative description of the stages of processing in central visual pathways implied a feed-forward system. Information was thought to be processed in a hierarchical manner through first-, second-, and third-order cells (Hubel and Wiesel 1962). More recent work has emphasized the possibility of intracortical processes that could involve feedback as in recurrent excitation models. (Ben-Yishai et al. 1995; Douglas et al. 1995; Somers et al. 1995). In the feed-forward process, the tuning of a postsynaptic cell is determined by pooling of information from many presynaptic cells. In the recurrent excitation model, feed-forward connections provide weak initial tuning that is then refined by feedback connections from neighboring cells, which also contribute broad based inhibition.
Either of these two mechanisms could produce a coarse-to-fine process such as the one we have described in the current study. Feed-forward processes could involve pooling of input from cells that have coarse-to-fine dynamics and pooling of input from cells with different spatial frequency content in which low-frequency latencies are shorter than those for high frequencies. Of course, both mechanisms may be involved. Although most of our current data may be accounted for by both feed-forward and feedback processes, this does not apply to the discrepancy between the monocular and binocular RF dynamics of simple cells. The feed-forward model cannot explain the discrepancy between the monocular RF and disparity-tuning size dynamics of binocular simple cells. Mathematically, this difference could be described as an exponent on the nonlinearity that increases with time. The greater the exponent, the smaller the disparity size becomes, without altering monocular RF size. Biologically, it is reasonable to speculate that there is disparity-tuned feedback from either complex cells or multiple simple cells. This results in monocular tuning that does not entirely predict disparity tuning as in the case of complex cells.
How does a coarse-to-fine process work? Low spatial frequency filters can respond to a wide range of disparities but with poor resolution. High spatial frequency filters have fine resolution but can only respond accurately to a limited range of disparities. Prior to the processing of disparity information, the system must solve a correspondence problem, i.e., it must select the correct right-left match so that the appropriate depth plane is identified. To do this, frequency and range information must be taken into account. An important theoretical proposal was put forward to address this issue. Information across spatial frequency scale may be combined so that low-frequency information constrains the range and this is followed by high-frequency resolution (Marr and Poggio 1979). The order of disparity information processing thus follows a coarse-to-fine sequence. The data we provide here are consistent with this notion. Our neurophysiological findings suggest a pooling across spatial frequency scale with a temporal bias from low to high frequencies that causes a coarse-to-fine process.
The original coarse-to-fine stereoscopic processing theory was followed by some refinements and variations (e.g., Nishihara 1984; Nomura 1993; Qian and Zhu 1997). A number of behavioral studies have been conducted to explore the relationships between low and high spatial frequency processing to determine if the data are compatible with the theory. There is clear psychophysical evidence that low spatial frequency information constrains processing on a fine scale (Rohaly and Wilson 1993; Wilson et al. 1991). On the other hand, some studies also suggest a reverse process by which high spatial frequency information is used to disambiguate that at low frequencies (Mallot et al. 1996; Smallman 1995; Smallman and MacLeod 1997). These processes could both occur by pooling across spatial frequency and averaging the result. In theory, this type of process can produce an unambiguous representation of disparity (Fleet et al. 1996). Disparity averaging across spatial scale has also been demonstrated psychophysically (Rohaly and Wilson 1994). Another approach to the idea of coarse-to-fine processing is to examine the temporal order of spatial frequency processing. In this case results show that low-frequency information is processed more rapidly than that of high values, i.e., there is a temporal coarse-to-fine mechanism (Glennerster 1996; Watt 1987). An additional relevant study shows that there is a transient stereopsis process that is temporally fast and consists of low spatial frequency information (Schor et al. 1998). This again is consistent with a coarse-to-fine process. It is important to point out that this type of mechanism may apply to other visual functions such as object recognition (Parker et al. 1997; Watt 1987). Considered together, the theoretical, behavioral, and neurophysiological studies point strongly to a processing system that begins with an approximation and ends with a fine-tuned percept.
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ACKNOWLEDGMENTS |
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TOPABSTRACTINTRODUCTIONMETHODSRESULTSDISCUSSIONACKNOWLEDGMENTSREFERENCES |
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GRANTS
This work was supported by research and CORE grants (EY-01175 and EY-03716) from the National Eye Institute.
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FOOTNOTES |
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Address for reprint requests and other requests: R. D. Freeman, 360 Minor Hall, Berkeley, CA 94720-2020 (E-mail: freeman{at}neurovision.berkeley.edu).
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REFERENCES |
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) is plotted at each correlation delay. Size decreases smoothly and extends beyond the optimal delay [thin regression line slope = –15.4 (%/10ms)].
, - - -) and RF envelope size parameters (
, P < 0.001, Bonferroni).
