Motion Perception Under Mesopic Vision

Journal of Vision (2016) 16(1):16, 1–15 1

Motion perception under mesopic vision Department of Psychology, Japan Women’s University, Kanagawa, Japan # Sanae Yoshimoto Japan Society for the Promotion of Science, Tokyo, Japan $ Faculty of Environment and Information Sciences, # Katsunori Okajima Yokohama National University, Kanagawa, Japan $ Department of Psychology, Japan Women’s University, # Tatsuto Takeuchi Kanagawa, Japan $

Mesopic and scotopic vision extend over an illuminance deals with this broad dynamic range by switching range of 106. The goal of the present study was to between two different types of photoreceptors: cones, determine the effect of decreasing light level on the which function at higher levels of illumination, and underlying motion mechanism that integrates rods, which function at lower levels. Photopic, mesopic, spatiotemporally separated motion signals. To and scotopic regions are defined according to whether accomplish this, we took advantage of the phenomenon cones operate alone, cones and rods operate together, of visual motion priming, in which the perceived or rods operate alone, respectively. In our daily lives, direction of a directionally ambiguous test stimulus is mesopic and scotopic vision extends over an illumina- influenced by the directional movement of a preceding tion range as wide as 106. Therefore, understanding the priming stimulus. After terminating a drifting priming effect of light levels on visual perception is scientifically stimulus, a 180 phase-shifted grating was presented as a 8 and practically crucial (Hess, 1990; Hess, Sharpe, & test stimulus. The priming and test stimuli were Nordby, 1990). separately presented to the central and peripheral retinas, respectively. The participants judged the Although rods project into all retinogeniculate perceived direction of this test stimulus at various light pathways, they primarily project into the magnocellular levels from photopic to scotopic levels. We found that lateral geniculate nucleus (LGN) layers (Purpura, the effects of motion priming disappeared over 1 log unit Kaplan, & Shapley, 1988; Zele & Cao, 2015). Because of mesopic light levels. When the test stimulus was motion-selective areas, such as the middle temporal presented before the offset of the priming stimulus to area, receive dominant inputs from the magnocellular compensate for the temporal delay in the rod pathway LGN layers, visual motion processing could be or when both stimuli were presented at the same selectively influenced by rod-based inputs (Hadjikhani location in the periphery, a motion-priming effect & Tootell, 2000; Maunsell, Nealey, & DePriest, 1990; appeared at mesopic light levels. These results suggest Maunsell & van Essen, 1983). In fact, various aspects of that different temporal characteristics between the cone human motion perception are known to change as light pathway and rod pathway disturb the function of the intensity decreases. Velocity perception (Gegenfurtner, putative motion mechanism responsible for the Mayser, & Sharpe, 2000; Hammett, Champion, spatiotemporal integration of motion signals, which Thompson, & Morland, 2007; Pritchard & Hammett, leads to specific modulation of motion perception over a 2012; Vaziri-Pashkam & Cavanagh, 2008), velocity wide range of mesopic vision. discrimination thresholds (Takeuchi & De Valois, 2000), short-range motion perception (Dawson & Di Lollo, 1990), complex-motion perception (Billino, Bremmer, & Gegenfurtner, 2008), biological motion Introduction perception (Billino et al., 2008; Grossman & Blake, 1999), perception of static-motion illusions (Hisakata & Ambient light levels may vary by a factor of up to Murakami, 2008), perception of interstimulus-interval 1011 in natural environments (Hood & Finkelstein, (ISI) reversal (Sheliga, Chen, FitzGibbon, & Miles, 1986; Stockman & Sharpe, 2006). The visual system 2006; Takeuchi & De Valois, 1997, 2009; Takeuchi, De

Citation: Yoshimoto, S., Okajima, K., & Takeuchi, T. (2016). Motion perception under mesopic vision. Journal of Vision, 16(1):16, 1–15, doi:10.1167/16.1.16.

doi: 10.1167/16.1.16 Received February 13, 2015; published January 27, 2016 ISSN 1534-7362

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Valois, & Motoyoshi, 2001), perception of two-stroke the spatially center-surround antagonistic motion- motion (Challinor & Mather, 2010; Mather & Challi- contrast sensing system (Allman, Miezin, & McGuin- nor, 2009), the coherent-motion threshold (Billino et ness, 1985; Born & Tootell, 1992; Eifuku & Wurtz, al., 2008; Lankheet, van Doorn, & van de Grind, 2002; 1998; Murakami & Shimojo, 1993; Tadin, Lappin, van de Grind, Koenderink, & van Doorn, 2000), Gilroy, & Blake, 2003). Antagonistic interactions moving texture segregation (Takeuchi, Yokosawa, & between the center and surround would induce a biased De Valois, 2004), and visual motion priming (Takeuchi, perception of motion direction for the directionally Tuladhar, & Yoshimoto, 2011; Yoshimoto & Takeuchi, ambiguous test stimulus in the peripheral visual field 2013; Yoshimoto, Uchida-Ota, & Takeuchi, 2014b) when the unidirectionally drifting stimulus is presented have all been shown to vary with the light level. in the central visual field. Since the priming and test Most of these studies have shown that motion stimuli were temporally separated in the visual motion- sensitivity decreases as light levels are reduced, priming display, the hypothesized mechanism should suggesting that changes in underlying temporal mech- be able to integrate motion signals separated by several anisms under low light levels affect motion perception. hundreds of milliseconds. However, Billino et al. (2008) measured the thresholds Then, we measured the effects of motion priming for the detection of biological motion under three under two lower light levels, presumably corresponding conditions of luminance corresponding to photopic, to mesopic (0.048 cd/m2) and scotopic levels (0.0048 cd/ mesopic, and scotopic light levels and found that m2). We found that under the mesopic level, the effects threshold was exclusively increased in the mesopic of motion priming completely disappeared. Under the condition, whereas the threshold under the scotopic scotopic level, however, negative motion priming was condition was identical to that under the photopic observed in a similar degree to that observed under the condition. The authors argued that in the mesopic photopic level (Yoshimoto & Takeuchi, 2013). As the condition, the mismatch of cone- and rod-mediated density of cones is higher in the central retina whereas velocity information led to impaired integration of the density of rods is higher in the periphery (Curcio, spatiotemporally separated motion signals, producing Sloan, Packer, Hendrickson, & Kalina, 1987; Oster- the largest threshold increase. Inspired by this finding, berg, 1935), our finding suggests that such a hypoth- we employed visual motion priming to examine a esized motion-contrast mechanism could not integrate motion mechanism under low light levels (Yoshimoto signals originated from cones at the center and rods at & Takeuchi, 2013). the periphery under mesopic vision. Visual motion priming is a phenomenon in which the We examined only one light level from the broad perceived direction of a directionally ambiguous test mesopic range extending over an illuminance range of stimulus is influenced by the direction of movement of 103–104 in our previous study (Yoshimoto & Takeuchi, the preceding priming stimulus. Examination of the 2013). However, motion information processing might effect of visual motion priming could reveal a be different even under mesopic vision because of the mechanism that integrates temporally separate motion complex nature of cone-rod interaction and the signals (Kanai & Verstraten, 2005; Pantle, Gallogly, & different amount of rod contribution across light levels Piehler, 2000; Pavan, Campana, Maniglia, & Casco, (Bloomfield & Dacheux, 2001; Cao, Pokorny, Smith, & 2010; Pinkus & Pantle, 1997). In these previous studies, Zele, 2008; Stockman & Sharpe, 2006; Zele & Cao, both priming and test stimuli were presented at the 2015; Zele, Maynard, Joyce, & Cao, 2014). Thus, in same location in the central visual field. Our previous Experiment 1 of the present study, we measured the work (Yoshimoto & Takeuchi, 2013) differed from strength of negative motion priming at various mesopic these studies by separately presenting the priming and light levels following our previous experimental method test stimuli in the central and peripheral visual fields in order to examine the effect of light levels on the under different light levels. The rationale for this integration of spatiotemporally separated visual inputs. manipulation was to create a situation in which the In our previous study (Yoshimoto & Takeuchi, induction of visual motion priming reflects a function 2013), we also found that negative priming reappeared of the underlying motion mechanism that integrates not at the mesopic light level when the test stimulus was only temporally, but also spatially, separate visual presented before the offset of the priming stimulus, inputs to induce motion perception. Our data showed causing a temporary overlap between the priming and that when the spatial distance between the priming and test stimuli. This indicates that the temporal delay in test stimuli was longer than 48, the test stimulus was the rod pathway at the periphery leads to the perceived as moving in the opposite direction of the disappearance of visual motion priming under mesopic priming stimulus (negative motion priming) under a vision. The temporal delay of the rod pathway relative high light level, presumably corresponding to a to the cone pathway has been estimated to be 70 ms at photopic level (48 cd/m2). We proposed that negative maximum (Sharpe & Stockman, 1999; Sharpe, Stock- priming could be induced by a mechanism similar to man, & MacLeod, 1989; Stockman & Sharpe, 2006).

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Figure 1. Schematic descriptions of stimuli in a single trial. The priming and test stimuli were presented to central and peripheral visual fields (VF), respectively. The spatial distance between the center of the priming stimulus and that of the test stimulus was set to 108. The black central cross (1.0831.08) was displayed to assist participants in maintaining fixation, while the grating was presented in the periphery. The participants were requested to judge whether the perceived direction of the test stimulus was rightward or leftward.

The temporal delay is reduced to 8–20 ms when the 3 768 pixels and 12-bit gray-level resolution. The state of adaptation is similar for the cones and rods monitor output was gamma-corrected with a Color- (Cao, Zele, & Pokorny, 2007; Sun, Pokorny, & Smith, CAL MKII colorimeter (Cambridge Research Systems 2001; Zele & Cao, 2015). In Experiment 1, we Ltd.). Neutral density filters (Kodak Wratten 2, manipulated the overlap time across a wide range of Edmund Optics Inc., Barrington, NJ) were placed in mesopic light levels to verify our previous observations. front of the monitor screen to obtain nine different photopic luminances (42, 3.0, 0.78, 0.21, 0.062, 0.022, 0.0065, 0.0024, and 0.00062 cd/m2); the luminances were checked by the same colorimeter. To convert Experiment 1 photopic troland (phot Td) to scotopic troland (scot Td), the spectrum of the monitor phosphors was Methods measured with a SpectroCAL MKII spectroradiometer (Cambridge Research Systems Ltd.). The participants Participants observed the display with the aid of a headrest. The Four participants (EA, SY, TT, and YI) with normal patterns were monocularly viewed with the right eye at or corrected-to-normal vision participated in the a distance of 57 cm. The room was darkened and experiments. SY and TT were coauthors of this study, shielded against external light. The fixation of the right and EA and YI were experienced in psychophysical eye for each participant was monitored using a experiments but na¨ıve to the experimental purpose. ViewPoint EyeTracker 220 fps USB system (Arrington This study was approved by the Research Ethics Research, Inc., Scottsdale, AZ) controlled by the same Committee of Japan Women’s University (Tokyo, PC during the entire experimental period. The sampling Japan) and was conducted according to the Declara- rate of this infrared video-based eye tracker was 220 tion of Helsinki. All participants provided written Hz. The same eye-tracking device was used to measure informed consent before the study began. the pupil diameter of each participant.

Apparatus Stimuli All stimuli were generated using Matlab (Math- Figure 1 illustrates a schematic description of the Works, Inc., Natick, MA) with the Psychophysics stimuli in a single trial. To allow comparisons with our Toolbox version 3.0 extension for PCs (Brainard, 1997; previous study (Yoshimoto & Takeuchi, 2013), we used Pelli, 1997) and were displayed on a 21-in color a similar stimulus. As the priming stimulus, a vertical monitor (GDM F520; Sony Corp., Tokyo, Japan) via drifting sine-wave grating was displayed in a rectan- VSG 2/5 (Cambridge Research Systems Ltd., Ro- gular window that measured 10.08 (width) 3 3.38 chester, UK) graphics system. The monitor temporal (height). The edges of the stimulus were tapered by a resolution was 120 Hz with a spatial resolution of 1024 Gaussian function with r ¼ 1.08. The spatial frequency

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Figure 2. Schematic representation of the timing relationship between the priming and test stimuli in Experiment 1. The horizontal and vertical axes represent time and luminance contrast, respectively. (A) There was no overlap time between the priming and test stimuli. (B) The overlap time between the priming and test stimuli varied from 8.3–83.3 ms. Both priming and test stimuli were presented during the overlap period, as illustrated.

of the stimulus was set to 0.5 c/8, which is well below stimulus by 1808. The total duration of the test stimulus the cut-off spatial frequency of about 6 c/8 under low was 667 ms. The priming and test stimuli were light levels (Hess et al., 1990). The stimulus was presented to the central (08 eccentricity) and peripheral presented on a uniform gray-colored background (108 eccentricity) upper visual fields, respectively (CIE1931; x ¼ 0.31, y ¼ 0.33) that had a similar (Figure 1). A black fixation cross (1.0831.08) was luminance to the space-averaged luminance of the sine- displayed to assist the participants in maintaining wave grating. The drift direction of the priming fixation while the test stimulus was presented in the stimulus was either rightward or leftward. Based on our periphery. previous study (Yoshimoto & Takeuchi, 2013), the In Experiment 1, the overlap time between the presentation duration and the velocity of the priming priming and test stimuli was varied in seven steps (0.0, stimulus were set to 167 ms and 6 8/s, respectively. The 8.3, 16.7, 25.0, 33.3, 41.7, and 83.3 ms). Figure 2 shows luminance contrast was set to two times that of the a schematic representation of the temporal relationship direction discrimination threshold, as described in the experimental procedure. between the priming and test stimuli. When the overlap An ambiguous test stimulus was generated by time was 0.0 ms, the test stimulus was presented shifting the phase of the grating by 1808, as in the immediately after the offset of the priming stimulus previous studies (Kanai & Verstraten, 2005; Pinkus & (Figure 2A). Pantle, 1997). The spatial frequency of the test stimulus was the same as that of the priming stimulus. To equate Experimental procedure the velocities of the priming and test stimuli, the phase of the test stimulus was shifted every 167 ms, which For each of three measurements described below, the corresponded to the time taken to shift the priming participants were dark-adapted for 30 min prior to the

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Figure 3. Individual and averaged photopic and scotopic retinal illuminances for the four participants obtained from the pupil diameter measurements. Scotopic retinal illuminance (log scot Td) is plotted as a function of the photopic retinal illuminance (log phot Td). Both vertical and horizontal error bars in the graph of averaged data represent 61 standard deviation (SD).

tasks. Each measurement started from the darkest this estimation, we assumed that the measured phot- adapting levels. opic retinal illuminances were categorized as one Pupil measurement: The retinal illuminances were photopic (2.8 log phot Td), seven mesopic (1.9, 1.3, computed from the participant pupil diameters, which 0.81, 0.33, 0.07, 0.57, and 0.99 log phot Td), and were measured under the nine luminances. A uniform one scotopic (1.6 log phot Td) light levels. Although space-averaged luminance blank field was presented for we used these categorizations as a reference in our 5 s while recording the pupil diameter. The participants Discussion, it is important to note that these represent were asked to focus on the center of the screen without a rough estimation, as we did not directly measure the blinking. Figure 3 presents both the individual and amount of cone/rod activation under different light averaged photopic and scotopic retinal illuminances for levels. the four participants. The averaged 5-s recording of the Contrast sensitivity measurement: For the main exper- pupil diameter range for the four participants was 3.9– iment (Figure 1), we equated the effective luminance 8.0 mm. The averaged nine retinal illuminances were contrast under different light levels and retinal eccen- approximated from 2.8 to 1.6 log phot Td. The tricities by setting the Michelson luminance contrast of conversion from phot Td to scot Td was calculated the sine-wave gratings to two times that of the direction from the spectral power distribution of the monitor discrimination threshold. For that, we measured the phosphors over a range of 380–780 nm. For the luminance contrast sensitivity for direction discrimi- computation, the CIE photopic V(k) modified by Judd nation of the drifting sine-wave gratings at each of the (1951) and Vos (1978) and the CIE (1951) scotopic nine light levels at 08 and 108 eccentricities. The V’(k) were used (Wyszecki & Stiles, 2000). The presentation duration of the drifting stimulus was 167 conversion factor for our monitor was 2.88. The ms, which was identical to that of the priming stimulus calculated log scot Tds are shown in Figure 3. used in the main experiment. The participants judged According to Hood and Finkelstein (1986) and the perceived drift direction of the stimulus with a Stockman and Sharpe (2006), the cone threshold and forced choice of two alternatives (rightward or rod saturation are approximately 1.4 and 1.9 log phot leftward) with no feedback. We applied a standard Td (or 1.0 and 2.3 log scot Td), respectively. Based on staircase algorithm that was designed to converge at a

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Figure 4. Individual and averaged data of the contrast thresholds obtained from direction discrimination measurements of the four participants. The percentage-contrast threshold is plotted as a function of the retinal illuminance (in log phot Td). Each curve represents data for the retinal eccentricity. The error bars in the graph of averaged data represent 61 SD.

79% correct level (Levitt, 1971) to estimate the Results threshold contrast. Further details of the experiment Contrast sensitivity measurement were described in Yoshimoto & Takeuchi (2013). When the stimulus was presented in the periphery, the Figure 4 shows the individual and averaged contrast participants were instructed to maintain their focus on thresholds for the direction discrimination of the moving sine-wave grating for the four participants at the cross pattern displayed. two eccentricities. The percentage-contrast threshold Direction judgment for estimating motion-priming ef- was plotted as a function of the photopic retinal fects: The priming stimulus was presented for 167 ms, illuminance. As shown in Figure 4, the contrast 500 ms after a beep (Figure 1). After terminating the thresholds depended on both the retinal illuminance priming stimulus, the directionally ambiguous test and the eccentricity. The contrast threshold increased stimulus was presented. The participants were asked to as the retinal illuminance decreased. At high light judge the perceived direction of the test stimulus by levels, the contrast thresholds at the center (08 pressing the appropriate arrow key. After a response, a eccentricity) were lower than those at the periphery (108 blank field was presented for 1 s during the intertrial eccentricity), whereas this tendency was reversed at low interval to reduce any effects from the previous trial. light levels. This result agrees with previous studies The participants were instructed to continuously view demonstrating that the motion sensitivity is higher at the fixation point throughout the trial. The partici- the periphery compared to the center under scotopic pants’ fixation was checked with an eye-tracking device vision (e.g., Hess et al., 1990). The transition of the during each trial. contrast threshold between the center and peripheral The direction judgment was conducted at an overlap fields was observed at 0.81 log phot Td under mesopic time of 0.0 ms and at an overlap time of 8.3–83.3 ms vision using averaged data. Similar results were during the separate sessions. For the overlap time of 0.0 observed for all four participants. These interpretations were supported by statistical ms, each session consisted of 32 trials: 16 trials for each analyses. We conducted a within-participant, two-way of the two priming stimulus directions which were analysis of variance (ANOVA) for the data. The presented randomly. Each participant completed a 2 generalized g G, which is suggested for analysis of session at each of the nine luminances. For the overlap within-participant designs (Bakeman, 2005; Olejnik & time of 8.3–83.3 ms, each session consisted of 96 trials: Algina, 2003), was used to estimate the effect size and eight trials for each of the six overlap times for the two interpreted according to Cohen’s recommendation of directions of the priming stimulus. Each participant 0.02 for a small effect, 0.13 for a medium effect, and completed two sessions at each of the nine luminances. 0.26 for a large effect (Cohen, 1988). The main effects The participants underwent at least 20 practice trials at of the retinal illuminance and eccentricity were 2 each light level prior to data acquisition. significant, F(8, 24) ¼ 113.2, p , 0.0001, g G ¼ 0.93 for

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Figure 5. Individual and averaged data for the four participants obtained from the direction judgment for estimating motion-priming effects. There was no overlap time between the priming and test stimuli. The percentage response to negative priming is plotted as a function of photopic retinal illuminance (in log phot Td). Error bars in the graph of averaged data represent 95% confidence intervals (CI).

2 retinal illuminance; F(1, 3) ¼ 43.19, p , 0.01, g G ¼ 0.44 Although there was some interparticipant variabili- for eccentricity. The interaction between the light level ty, these results were essentially consistent across the and eccentricity was also significant, F(8, 24) ¼ 9.56, p participants. Thus, the averaged data were used in our 2 , 0.0001, g G ¼ 0.52. analyses. As shown in Figure 5, we found that the effect In the following experiments, we set the luminance of motion priming greatly depended on the retinal contrast of the gratings to be twice the measured illuminance. A within-participant, one-way ANOVA threshold for each participant. revealed that the effect of retinal illuminance was 2 significant, F(8, 24) ¼ 40.63, p , 0.001, g G ¼ 0.91. Under photopic conditions (2.8 log phot Td), negative Direction judgment for estimating motion-priming priming was observed in the majority of trials, which effects replicated our previous results (Yoshimoto and Take- The number of trials in which participants looked uchi, 2013). At higher mesopic light levels (1.9 and 1.3 more than 1.58 away from the fixation point through- log phot Td), negative priming was reported in more than 90% of the trials. However, at 0.81 log phot Td, out a single trial was less than 1%. Thus, excluding data the percentage of negative motion priming was from these trials did not change our results and approximately 50% (95% CI of [43.35%, 61.4%]), which conclusions. Therefore, we used the data from all trials indicates that the priming effect totally disappeared and for the subsequent analysis. Figure 5 shows the did not recover for retinal illuminances of 0.33 (95% CI individual and averaged data for the four participants of [45.7%, 63.7%]) and 0.07 log phot Td (95% CI of when there was no overlap time between the priming [44.9%, 62.9%]). Thus, neither positive nor negative and test stimuli (Figure 2A). The percentage response priming was dominant for an approximate 1 log unit to the negative motion priming is plotted as a function range under mesopic conditions. The priming effect of the retinal illuminance (in log phot Td). When more gradually reappeared from mesopic (0.57 log phot Td) than 50% of the responses represented negative motion to scotopic (1.6 log phot Td) levels, where negative priming, the participants reported that the perceived priming became dominant. A reversal of the contrast direction of the test stimulus was opposite to that of the threshold for the central and peripheral retinas priming stimulus in the majority of the trials. When occurred at 0.81 log phot Td (Figure 4), which was fewer than 50% of the responses were scored as coincident with the retinal illuminance where the negative priming, the participants reported that the priming effect disappeared in Figure 5. motion was in the same direction as the priming Figure 6 shows the individual and averaged data for stimulus (the so-called positive motion priming) in the the four participants when the priming and test stimuli majority of trials. temporally overlapped. The data at the 0.0-ms overlap

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Figure 6. Individual and averaged data for the four participants. The percentage response to negative priming is plotted as a function of photopic retinal illuminance (in log phot Td). Each curve represents data for the overlap times between the priming and test stimuli (0.0–83.3 ms). The pale blue curves (overlap time of 0.0 ms) are replots of data from Figure 5. The red arrows in the averaged data graph represent mesopic conditions at which no priming effect was observed at the overlap time of 0.0 ms (0.81, 0.33, and 0.07 log phot Td). Error bars in the graph of averaged data represent 61 standard error of the mean.

time (pale blue curves) were replotted from Figure 5. different between 0.81 and 0.07 log phot Td (indicated Although there was some interparticipant variability, by the rightmost and leftmost red arrows, respectively). these results were essentially consistent across the Thus, we selected these data (0.81, 0.33, and 0.07 log participants. Thus, the averaged data were used in our phot Td) and replotted the percentage of negative analyses. priming as a function of the overlap time in Figure 7. As the overlap time between the priming and test The overlap time required to induce the conspicuous stimuli increased, the function changed from a U- to a negative priming at 0.81 log phot Td was shorter than flat-shape (Figure 6). In the mesopic conditions the overlap time required at 0.33 or 0.07 log phot Td. indicated by red arrows (0.81, 0.33, and 0.07 log phot At the overlap time of 8.3 ms, negative priming was Td), the effect of increasing the overlap time was clear. observed in more than 80% of trials at 0.81 log phot In addition, negative priming was induced by increas- ing the overlap time at lower mesopic levels (0.57 and 0.99 log phot Td). In contrast, at the two highest mesopic levels (1.9 and 1.3 log phot Td) as well as under photopic (2.8 log phot Td) and scotopic (1.6 log phot Td) conditions, negative priming was dominant irrespective of the overlap time. At overlap times longer than 25.0 ms, negative priming was observed in more than 70% of trials irrespective of retinal illuminance. Statistical analyses support these interpretations. A within-participant, two-way ANOVA was conducted on the average data depicted in Figure 6. The main effects of the retinal illuminance and overlap time were 2 significant, F(8, 24) ¼ 17.05, p , 0.0001, g G ¼ 0.64 for 2 retinal illuminance; F(6, 18) ¼ 34.80, p , 0.0001, g G ¼ 0.50 for overlap time. The interaction between retinal illuminance and overlap time was also significant, F(48, Figure 7. The percentage response to negative priming was 2 144) ¼ 6.50, p , 0.0001, g G ¼ 0.55. plotted as a function of the overlap time between the priming A close examination of Figure 6 revealed that the and test stimuli. Each curve represents data for different retinal shapes of the functions for the reappearance of negative illuminances, which are indicated by red arrows in Figure 6 priming which depended on the overlap time were (0.81, 0.33, and 0.07 log phot Td). Error bars represent 95% CI.

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Td, whereas the priming effect was not observed at 0.33 antagonistic motion-contrast sensing system is respon- (95% CI of [48.06%, 65.99%]) and 0.07 log phot Td, sible for the motion-priming effect. Based on this, one (95% CI of [47.27%, 65.23%]). possible explanation for the disappearance of negative A within-participant, two-way ANOVA followed by priming at some mesopic light levels is that the Tukey’s posthoc test was conducted for multiple temporal delay in the rod pathway fails to provide comparisons. The main effects of the retinal illumi- visual inputs within the given time frame that would nance and overlap time were significant, F(2, 6) ¼ 5.27, allow such a mechanism to operate. The assumed 2 p , 0.05, g G ¼ 0.30 for retinal illuminance; F(6, 18) ¼ temporal delay would be shorter at higher light levels 2 38.76, p , 0.0001, g G ¼ 0.81 for overlap time. The (Figure 7), which may correspond to the known interaction between the retinal illuminance and the temporal characteristics of the rod pathway (Sharpe et overlap time was also significant, F(12, 36) ¼ 4.02, p , al., 1989; Zele & Cao, 2015). 2 0.001, g G ¼ 0.31. The Tukey’s test revealed significant The negative priming was observed at the two differences in the percentage of negative priming highest mesopic light levels (1.3 and 1.9 log phot Td) between 0.81 and 0.07 log phot Td at overlap times of and at mesopic light levels lower than 0.57 log phot 8.3 (q ¼ 7.70, p , 0.0001), 16.7 (q ¼ 6.49, p , 0.001), Td. This may be because that the primarily operating 25.0 (q ¼ 5.75, p , 0.001), and 33.3 ms (q ¼ 3.78, p , system is the cone system at the higher mesopic levels 0.05). The Tukey’s test also showed significant differ- and it is the rod system at the lower mesopic levels ences between 0.81 and 0.33 log phot Td at overlap across the retina, which did not induce temporal times of 8.3 (q ¼ 7.50, p , 0.0001) and 16.7 ms (q ¼ conflict between the cone and rod pathways. We 6.49, p , 0.001). No significant difference was found examined this conjecture in the next experiment. between 0.33 and 0.07 log phot Td.

Discussion Experiment 2

The changes of contrast thresholds and motion In Experiment 2, we examined whether motion priming along with decrements in retinal illuminance priming would occur under mesopic conditions by were quite different (Figures 4 and 5). The contrast presenting both the priming and test stimuli at the same thresholds gradually decreased for both the central and peripheral location. Because no temporal conflict peripheral fields, whereas negative motion priming would arise between the cone and rod pathways with showed a sudden disappearance and gradual recovery this manipulation, we predicted that the motion- as the retinal illuminance decreased on a log scale. This priming effect would be observed regardless of retinal difference suggests that the underlying motion mecha- illuminance. nism responsible for the direction discrimination of the moving stimulus is different from that for the induction of negative motion priming. In the direction discrim- Methods ination task, a motion detector which is mostly tuned to a given stimulus would determine the luminance Here, we presented the test stimulus at a 108 contrast threshold (e.g., Watson, 1986). On the other eccentricity, similar to Experiment 1. However, we also hand, negative motion priming observed at a supra- presented the priming stimulus at the same peripheral threshold luminance contrast would reflect the inte- location as that of the test stimulus, not at the central gration of the outputs from various motion detectors retina, as in Experiment 1. To examine the effect of the that detect spatiotemporally separated priming and test rod system on the motion-priming effect, the luminance stimuli (Pinkus & Pantle, 1997; Yoshimoto & Take- level was varied from mesopic (3.0 cd/m2 or 1.9 log uchi, 2013). phot Td) to scotopic light levels (0.00062 cd/m2 or 1.6 In Experiment 1, we found that the motion-priming log phot Td) in eight steps. There was no overlap time effect disappeared around 1 log unit of the mesopic between the priming and test stimuli. All other range, approximately from 0.8 to 0.1 log phot Td parameters were the same as in Experiment 1. (Figure 5). In light of the distinct distributions of cones Each session consisted of 32 trials: 16 trials for each and rods in the retina (Curcio et al., 1987; Osterberg, of the two directions of the priming stimulus, which 1935), we hypothesize that at this mesopic range, the were presented randomly. Each participant completed motion signals from both the cone and rod systems in one session at each of the eight retinal illuminances the center and the motion signals mainly from the rod (Figure 3), starting from the darkest adaptation level. system in the periphery could not integrate well. As In each session, the light level was fixed. The same described in the Introduction, we speculate that a participants that performed Experiment 1 participated mechanism similar to the spatially center-surround in Experiment 2.

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Figure 8. Individual and averaged data for the four participants in Experiment 2. The percentage response to negative priming is plotted as a function of photopic retinal illuminance (in log phot Td). Each curve represents data for the eccentricities of the priming and test stimuli. ‘‘108–108’’ denotes that both stimuli were presented in the periphery (108 eccentricity). The pale blue curves (08–108) are replots of data from Figure 5 in which the priming stimulus was presented to the central retina (08 eccentricity), and the test stimulus was presented to the periphery (108 eccentricity). Error bars in the graph of averaged data represent 95% CI.

Results and discussion mesopic light levels as well as at the photopic and scotopic levels (Figure 5). Figure 8 shows the individual and averaged data for 2. In the mesopic range where the priming effect the four participants. The Experiment 1 data for the disappeared, the frequency of reporting negative priming and test stimuli, which were presented sepa- priming gradually increased with an increase in the rately to the central and peripheral fields, was replotted overlap time between the priming and test stimuli from Figure 5. Although there was some interpartici- (Figure 6). The duration of the overlap time that pant variability, the results were relatively consistent induced negative priming was shorter at higher across participants. Thus, the averaged data were used mesopic levels (Figure 7). in our analyses. In contrast to Experiment 1, we found 3. When the priming and test stimuli were presented at that negative priming was observed in more than 90% the same peripheral location, negative priming was of trials, irrespective of retinal illuminance. A within- dominant irrespective of retinal illuminance (Figure participant, one-way ANOVA for the averaged data 8). revealed that there was no significant effect of retinal The relative amount of rod contribution should have illuminance, F(7, 21) ¼ 1.77, n.s. This result supports been different across the retina at the mesopic light our hypothesis that the temporal delay in the rod levels, since both the cones and rods are active in the pathway disturbs the spatiotemporal integration of the central retina, while the rods are mainly active in the motion signals during mesopic vision. peripheral retina (Raphael & MacLeod, 2011). Our results indicate that the spatiotemporally separate motion signals were not well integrated when the General discussion motion signals were processed via the central and peripheral retinas. This conclusion is in accordance with the findings reported by Billino et al. (2008), which Summary and conclusion demonstrated that the simultaneous activity of the cones and rods may exert a detrimental effect on We summarize the findings of our study below: motion integration, resulting in a mesopic-specific threshold elevation. The mesopic light level of 0.285 cd/ 1. The effect of motion priming disappeared at the m2 (or about 0.9 log phot Td) used in their study is approximately 1 log unit range of mesopic light similar to our highest mesopic light range, where we levels (0.8 to 0.1 log phot Td). Negative motion observed a disappearance of motion priming (from priming was observed at the higher and lower approximately 0.8 to 0.1 log phot Td).

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We also demonstrated that the motion-priming nance imaging, Hadjikhani and Tootell (2000) dem- effect reappeared when the overlap time of the priming onstrated that the signals from the cone and rod and test stimuli was increased (Figure 6) or when the pathways are combined at the level of V1. By applying temporal conflict between the cone and rod pathways repetitive transcranial magnetic stimulation, Campana was removed (Figure 8). These results indicate that the et al. (2011) identified the involvement of V1 when incomplete spatiotemporal integration of motion in- negative motion priming was induced. Taken together, formation may lead to the disappearance of motion our findings indicate that under mesopic vision, the priming. This is presumably due to a maximum delay failure to provide visual input to the motion mechanism of approximately 20 ms in the rod pathway from the in V1 at the appropriate timing results in incomplete cone pathway in the peripheral retina (Figure 7). motion integration. Although further studies are needed to confirm this, the estimation of the rod delay obtained here is consistent with the current estimations of the cone-rod latency Effect of presentation duration on visual motion difference in similar states of light adaptation (Cao et priming al., 2007; Sun et al., 2001; Zele & Cao, 2015). Previous studies have shown that when both priming and test stimuli are presented at the same location in Underlying mechanism for negative motion the fovea, a priming stimulus as short as 300 ms induces priming positive priming in which the test stimulus is perceived to drift in the same direction as the priming stimulus We hypothesize that a mechanism similar to the (Kanai & Verstraten, 2005; Pantle et al., 2000). In spatially center-surround antagonistic motion-contrast contrast, we found that when both priming and test sensing system is a candidate for the underlying stimuli are presented at the same peripheral retinal mechanism for negative motion priming. In this location, negative priming is induced by the 167 ms section, we suggest alternative explanations. One is priming stimulus (Figure 8). This finding suggests that based on the studies examining a related phenomenon the effect of the priming stimulus presentation duration called the phantom or remote motion aftereffect— depends on the retinal eccentricity of moving patterns: MAE (von Gru¨nau & Dube´, 1992; Snowden & Milne, The duration that switches from positive to negative 1997). In this stimulus configuration, the test stimulus priming effect may decrease as the retinal eccentricity was presented at a different spatial location after the increases. adapting moving stimulus was presented for 30–60 s. We speculate that the multisystem for motion The test stimulus was then perceived to move in the perception is responsible for this dependency on retinal opposite direction of the adapting stimulus. Our data eccentricity. Our previous studies (Takeuchi et al., indicate that a similar phenomenon could be induced 2011; Yoshimoto & Takeuchi, 2013; Yoshimoto, by a short-term adaptation (167 ms) to the moving Uchida-Ota, & Takeuchi, 2014a) suggested that nega- stimulus, if a dynamic (i.e., temporally varying) test tive priming is induced by an energy-based, direction- stimulus is used. Snowden and Milne (1997) demon- ally selective, first-order motion mechanism (Adelson & strated that the phantom MAE was observed in an area Bergen, 1985; van Santen & Sperling, 1985; Watson & with a 58 diameter and concluded that motion detectors Ahumada, 1985), which is shown to be sensitive to tuned to wide-field motions induced the phantom moving stimuli with low-contrast and presented in the MAE. This model is a feasible candidate for the periphery (Ashida, Seiffert, & Osaka, 2001; Chubb & underlying mechanism of the negative priming effect Sperling, 1989; Dosher, Landy, & Sperling, 1989; during photopic vision reported in this study. Another Edwards & Nishida, 2004; Lorenceau & Shiffrar, 1992; candidate is a mechanism that assumes a long-range Lorenceau, Shiffrar, Wells, & Castet, 1993; Lu & suppression between spatiotemporally separated loca- Sperling, 1995, 2001; Smith, Hess, & Baker, 1994; tions. Such a mechanism has been proposed for the Solomon & Sperling, 1994, Sperling, 1989; Sun, Chubb, spatial domain (Polat & Sagi, 1993), but recent studies & Sperling, 2014, 2015; Takeuchi & De Valois, 1997, have demonstrated a long-range interaction in the 2009; Weiss, Simoncelli, & Adelson, 2002; Yo & spatiotemporal domain (Yeshurun, Rashal, & Tkacz- Wilson, 1992). In the current study, the prominent Domb, 2015). negative priming disappeared when the test stimulus Although further studies are needed to identify the was presented in the periphery at two times the underlying mechanism of visual motion priming, our threshold contrast (i.e., low contrast) during mesopic results indicate that the spatiotemporal characteristics vision (Figure 5). This suggests that the function of the in the early visual pathway, presumably at the retinal first-order motion mechanism can be influenced under level, critically influence the later stages of visual mesopic vision. Further studies are required to clarify motion processing. Using functional magnetic reso- the relationship between the light level-dependency of

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