Article

Perception
2023, Vol. 52(3) 195–212
Categorical encoding of moving © The Author(s) 2022
Article reuse guidelines:
colors during location tracking sagepub.com/journals-permissions
DOI: 10.1177/03010066221147120
journals.sagepub.com/home/pec

Mengdan Sun
Soochow University, China

Xiaoyang Xin
Zhejiang University, China

Haojiang Ying
Soochow University, China

Luming Hu
Beijing Normal University, China

Xuemin Zhang
Beijing Normal University, China

Abstract
Categorical perception (CP) describes our tendency to perceive the visual world in a categorical
manner, suggesting that high-level cognition may affect perception. While most studies are con-
ducted in static visual scenes, Sun and colleagues found CP effects of color in multiple object track-
ing (MOT). This study used functional magnetic resonance imaging to investigate the neural
mechanism behind the categorical effects of color in MOT. Categorical effects were associated
with activities in a broad range of brain regions, including both the ventral (V4, middle temporal
gyrus) and dorsal pathways (MT + /V5, inferior parietal lobule) of feature processing, as well as
frontal regions (middle frontal gyrus, medial superior frontal gyrus). We proposed that these
regions are hierarchically organized and responsible for distinct functions. The color-selective
V4 encodes color categories, making cross-category colors more discriminable than within-cat-
egory colors. Meanwhile, the language and/or semantic regions encode the verbal information
of the colors. Both visual and nonvisual codes of color categories then modulate the activities
of motion-sensitive MT + areas and frontal areas responsible for attentional processes.

Corresponding author:
Xuemin Zhang, Beijing Key Lab of Applied Experimental Psychology, Faculty of Psychology, Beijing Normal University,
Beijing 100875, China.
Email: xmzhang@bnu.edu.cn
196 Perception 52(3)

Keywords
color category, perception, language, multiple object tracking, fMRI
Date Received: 29 July 2022; accepted: 7 December 2022

While we perceive colors continuously in terms of hue, lightness, and saturation, we only use a
handful of discrete color categories (e.g., blue, green) in the language system. The categorical per-
ception (CP) effect of color has been repetitively shown in various tasks that color stimuli from dif-
ferent categories are more discriminable than those from the same category (e.g., Bornstein &
Korda, 1984), traditionally considered evidence of language-perception interaction (for review,
see Siuda-Krzywicka et al., 2019; Witzel, 2019). While most CP studies were conducted in
static visual scenes, Sun and colleagues (Sun et al., 2018, 2020; Sun & Zhang, 2022) previously
explored the effects of color categories on tracking performance using the multiple object tracking
(MOT) paradigm in which observers were asked to track the locations of multiple moving targets
among. These MOT studies manipulated the categorical relationship between the surface colors of
the multiple moving objects. They implicitly probed the effects of color categories in visual pro-
cessing through location tracking instead of explicit judgment based on color appearance.
Categorical effects were found that tracking within-category targets was easier than tracking cross-
category targets when the targets’ categorical relationship was varied (Sun et al., 2020). Color cat-
egories did not influence tracking performance when the categorical relationship between the
targets and distractors was manipulated (Sun et al., 2018). In contrast to a cross-category advantage
found in most previous CP studies, the CP effects in MOT (Sun et al., 2020) are manifested as a
cross-category disadvantage for the first time. Furthermore, Sun and Zhang (2022) found that
the CP effects in tracking can be partly disrupted by a secondary verbal interference task, suggesting
an online use of language (Sun & Zhang, 2022). Interestingly, the interference effect only emerged
in the cross-category trials, which implies that the online use of color names without interference
affects tracking in a way that makes tracking cross-category targets harder. In other words, the CP
effects emerge automatically in MOT even when encoding color categories has a detrimental
impact on tracking performance. These findings suggested that color categories affect not only
direct discrimination of the color stimuli (e.g., Winawer et al., 2007) but also how we track and
organize multiple color objects in an implicit way. However, the CP effects observed in MOT
can be related to the CP effects in color discrimination. Performing MOT requires successful seg-
mentation between the targets and the distractors. When the targets are less discriminable from one
another in feature, they are easier to be grouped and tracked simultaneously (Endress et al., 2017;
Erlikhman et al., 2013).
Following the previous MOT studies that observed CP effects (Sun et al., 2020; Sun & Zhang,
2022), the current study used functional magnetic resonance imaging (fMRI) to investigate the
neural mechanisms behind CP effects in MOT. Regarding the representation of color categories
in the human brain, previous evidence remains mixed (Bird et al., 2014; Brouwer & Heeger,
2013; Liu et al., 2019; Persichetti et al., 2015; Siok et al., 2009; Tan et al., 2008; Yang et al.,
2016). The reported brain regions include the visual cortex (Brouwer & Heeger, 2013;
Persichetti et al., 2015; Siok et al., 2009; Yang et al., 2016), language-related networks (Siok
et al., 2009; Tan et al., 2008), and frontal areas subserving domain-general categorization (Bird
et al., 2014; Liu et al., 2019; Persichetti et al., 2015). Some studies revealed that the extrastriate
cortex is category-selective, such as V4 (Brouwer & Heeger, 2013; Persichetti et al., 2015; Siok
et al., 2009). For example, Brouwer and Heeger (2013) found that within-category colors
showed categorical clustering in V4 and VO1 when participants actively categorized color
Sun et al. 197

stimuli but not when they passively observed them. Using the fMRI adaptation paradigm,
Persichetti et al. (2015) found that color categories failed to mediate the adaptation effect of
BOLD signals in the visual cortex. However, they observed increased BOLD responses to
stimuli near the category boundary in visual areas, especially during the categorization task. The
language processing areas were also found to be engaged in perceptual discrimination of colored
stimuli (Siok et al., 2009; Tan et al., 2008). Tan et al. (2008) observed different activity between
easy-to-name and hard-to-name colors in the left posterior STG and IPL that subserve
color-to-word mapping. Beyond the classic language-perception debate, some studies observed
encoding of color categories in the prefrontal cortex associated with domain-general categorization
(Bird et al., 2014; Liu et al., 2019; Persichetti et al., 2015), including the middle frontal gyrus
(MFG), precentral sulcus, inferior frontal gyrus, superior frontal sulcus, intraparietal sulcus, and
pre-supplementary motor area.
According to these findings, it is evident that the type of visual tasks can modulate the stage
where the categorical effects occur, as they engage different levels of cognitive processes. Thus,
it is interesting whether the categorical effects observed in tracking are due to categorization in
the perceptual stage or higher-level stages. In this study, participants were required to keep track
of six moving objects (i.e., targets). Among the targets, three shared one surface hue, and the
other three shared another surface hue. Importantly, we manipulated the categorical relationship
(within-category vs. cross-category) and hue distances (large vs. small) between these two hues.
Unlike the previous study (Sun et al., 2020), no distractors were included in the current study.
The rationale for not including distractors is twofold. First, Sun et al. (2020) revealed two kinds
of CP effects: target CP effects (i.e., color categories of the targets influence tracking performance)
and distractor CP effects (i.e., color categories of the distractors influence tracking performance).
Although the distractor CP effects are task-dependent and not as robust as the target CP effects,
it is suggested that the categorical relationship of the distractors is encoded to some extent
during tracking. The current study intends to focus on the target CP effects and minimize the influ-
ence of the distractors’ categorical relationship. We ensure that all colors are from the same cat-
egory in the within-category condition (distractors must be cross-category if included). In
addition, excluding distractors means fewer stimulus samples, which reduces the problem caused
by perceptual inequality. However, the exclusion of distractors posed difficulty testing the tracking
performance, which is commonly indexed by the proportion of correctly tracked targets. Under this
circumstance, we included a probe detection task in which a probe stimulus would appear randomly
around one of the moving targets at random moments to ensure participants were tracking the loca-
tion of the targets. Even though there was a high probability of correctly detecting the probe while
participants were not tracking the moving targets, the probe detection performance would also be
affected by the surface feature of the targets. Thus, if we observed differences in probe detection
performance between conditions, it means that the participants were truly performing the location
tracking task. Otherwise, the detection accuracy would be homogenous across conditions.
In summary, color category (within-category, cross-category) is the primary independent vari-
able in our study. We were interested in how color categories influence both tracking performance
and brain activities. In addition, we included hue distance as an independent variable, which allows
us to compare the brain activities induced by categorical differences with those by perceptual dif-
ferences (i.e., hue) and determine the unique neural circuits responsible for CP effects in MOT.
Based on the CP effects observed in several previous studies (Sun et al., 2020; Sun & Zhang,
2022), our first prediction is that color categories will influence the tracking (i.e., probe detection)
performance that participants can perform better with within-category targets than cross-category
ones. Concerning the brain regions involved in CP effects in MOT, we intend to reveal neural cir-
cuits supporting two distinct functions: (1) encoding color categories and (2) involved in the effect
of color categories on attentive tracking. The candidate brain areas for categorical encoding are
198 Perception 52(3)

visual cortex (Brouwer & Heeger, 2013; Persichetti et al., 2015; Siok et al., 2009; Yang et al.,
2016), language-related (Siok et al., 2009; Tan et al., 2008), and frontal areas (Bird et al., 2014).
With regard to the effect of color categories on tracking, one plausible hypothesis is that the
shared categorical feature (e.g., green) could be utilized to group targets into a virtual object
(Endress et al., 2017; Lupyan, 2008; Wei et al., 2017), while different categorical features (e.g.,
green and blue) disrupt this kind of grouping. If the grouping hypothesis is accurate, we expect
that the brain regions associated with shape and object processing (e.g., lateral occipital cortex,
LO) should be involved. The other hypothesis is that color categories influence identity-location
binding during tracking, which predicts increased activities in the lateral frontal cortex for the cross-
category than within-category stimuli. The lateral prefrontal cortices were suggested to be the
central “hubs” of the object-identity-tracking network (Nummenmaa et al., 2017).

Materials and Methods

Participants
Two independent groups of participants were recruited in this study, with one group (6 participants,
age: 20.83 ± 3.76 years) in the preliminary behavioral session and the other group (17 participants,
age: 20.82 ± 3.07 years) in the formal fMRI session. All participants are healthy right-handed
undergraduate and graduate students with normal (corrected-to-normal) vision and color vision.
None of them had any neurological or psychological disorders. All participants gave written
consent before the experiment and received payment after.

Stimuli
The color stimuli used for the tracking task were chosen from CIELAB space (L∗ = 70, a radius of
29), which only varied in hue (Figure 1A). The green-blue boundary is 192.1° (SD = 6.9°), deter-
mined according to a preliminary color naming task outside the MRI scanning room. In this prelim-
inary naming task, 11 colors were used (140–240°, step size: 10°). On each trial, one color disk
appeared in the center of the screen, and the participants were required to indicate whether the
color was green or blue by pressing “F” or “J”. The proportion of “blue” responses in the color
naming task was fitted with a psychometric function defined as 1/ (1 + exp ( − (x − α)/β), where
α is the threshold (estimated value at which “blue” would be reported half of the time), that is,
the pre-determined green–blue boundary in our study. Note that the participants in the preliminary
color naming task were not recruited in the following scanning session.
There were four conditions (Figure 1A): Large Within (LW), Large Cross (LC), Small Within
(SW), and Small Cross (SC). Under the Within conditions (i.e., LW, SW), the objects shared two
different green colors. Under the Cross conditions (i.e., LC, SC), the objects shared one green color
and one blue color. The distance between two hues was 40° under the Large conditions, and it was
20° under the Small conditions.
During scanning, the stimuli were projected onto a screen placed at the back of the magnet bore
from a video projector. Participants viewed this screen (48 cm × 36 cm) through a mirror, which
was at a distance of approximately 110.5 cm from their eyes. So the effective size of the display
was 24.5° by 18.4°. All stimuli were presented inside a rectangle area with a gray background.
The moving stimuli are six colored disks (0.5° radius). The screen was calibrated using Minolta
Konica CS-150 colorimeter (https://sensing.konicaminolta.us). The xyY value of the white point
(x = 0.4106, y = 0.4464, Y = 49.74) and the screen primaries (R: x = 0.6342, y = 0.3457, Y =
8.66; G: x = 0.3895, y = 0.5740, Y = 25.40; B: x = 0.1805, y = 0.1353, Y = 2.41) were measured.
The input–output value of each channel was also measured to determine the gamma curve. This
Sun et al. 199

Figure 1. Stimuli, procedure, and ROI definition. (A) Illustration of color stimuli used in four conditions
(LW: Large Within; LC: Large Cross; SW: Small Within; SC: Small Cross). (B) Sample of a trial. T1: fixation
stage (500 ms); T2: all color disks started to move randomly and independently; T3: a probe was flashed
around one of the objects during a random period between 2 and 5 s after movement; T4: response stage.
(C) Brain regions showing positive task effects in the magnitude of hemodynamic response function (HRF).
(D) Two functional (left MT + and right MT + ) and four structural (left V4, right V4, left V4a, right V4a) ROIs
were defined as spheres with a radius of 6 mm centered at the peak voxels in the significant clusters. The
figure only shows the right hemisphere for illustration.

information was used to determine the appropriate RGB values for each color stimulus suggested
by Brainard et al. (2002).

Procedure
The tracking task adopted a block design. Each trial begins with a white fixation on a gray back-
ground (L∗ = 50 in CIELAB color space) for 500 ms (Figure 1B). Then six disks were presented
in random locations inside a rectangle area with a white border and started to move independently
and randomly at an average speed of 8°/s. Participants were asked to keep track of them and
informed that a probe would appear around the moving disks. In other words, they should attend
to these six disks simultaneously. To ensure that the participants were performing the tracking
task, a white rectangle, as the probe, flashed around one of the disks during a random period
between 2 and 5 s after movement on half of the trials. The participants were required to press a
button once they detected the probe. After 5.5 s, all the objects stopped moving (please see the
demo in Supplemental Material). Each trial lasted for 6 s in total.
Four conditions were presented in pseudo-random sequences ([A C B D B C A D], [D A B C D
C B A], [D C A B A C D B]), forming three runs. The sequence was arranged so that the types of
condition-switching (e.g., A-B) within each sequence were heterogeneous. Each run includes eight
blocks of five trials of one condition, and therefore, there were 40 trials in total in each run.
200 Perception 52(3)

Participants rested and viewed a gray background for 2 s after each block, except for the last block
in each scan. There was a 1 min rest after each of the first two functional scans. Before scanning, the
participants were trained for 5 min outside the scanner to understand the tracking process fully.

Image Acquisition
Scanning was performed in a 3T MRI scanner (Siemens Magnetom Trio, A Tim System). A stand-
ard 12-channel head coil was used for coverage of the entire brain. The high-resolution
T1-weighted MPRAGE anatomical images (TR = 2300 ms, TE = 2.86 ms, flip angle = 9°, 144
slices, matrix size = 256 × 256, voxel size = 1 × 1 × 1.33 mm3) were acquired for each participant
before the tracking task. Functional images were acquired with a gradient-echo single-shot echo-
planar sequence (TR = 2000 ms, TE = 30 ms, flip angle = 90°, 33 slices, inter-leaved slice order,
matrix size = 64 × 64, FOV = 200 × 200 mm, voxel size = 3.125 × 3.125 × 4.025 mm3).

Data Analysis
Category Membership
Since the assignment of the within- or cross-category condition in tracking was based on the pre-
liminary naming task with independent participants, we first assessed whether the assignment was
appropriate for the participants who performed the tracking task. According to the results of another
study (Sun et al., 2021) where the same group was recruited in an active color naming task, we
found that the average subjective green-blue boundary was 200.74° (SD = 6.70°). The Small
Cross condition (180°, 200°) defined based on the mean subjective boundary (192.1° ± 6.9°) of
the participants in the preliminary task was not a genuine cross-category condition at the group
level. We included only the Large Within and Large Cross conditions when analyzing category
effects. Note that we conducted the current study and then the color naming study (Sun et al.,
2021) on the same day with the same group of subjects. Thus, we were not able to use the boundary
data (Sun et al., 2021) to select color samples for the tracking task.
Furthermore, we reanalyzed the behavioral data in Sun et al. (2021) and defined category dis-
tance as a continuous index of subjective categorical differences for color stimuli in cross condi-
tions. For each participant, the categorical distance for a pair of color stimuli was denoted by
their difference in the proportion of ‘blue’ responses in the behavior color naming task. For
instance, if a participant’s proportion of ‘blue’ response was 0.2 for Hue 170° and 1.0 for Hue
210°, then the categorical distance of this participant in the Large Cross condition was 0.8. This
measure was used for further analyses.

fMRI Analyses
Preprocessing and statistical analyses of fMRI data were performed using SPM12 (Wellcome
Department of Imaging Neuroscience, University College London, UK, http://www.fil.ion.ucl.ac.
uk/spm) for each run. Preprocessing the functional data included slicing timing, realignment, cor-
egistration, spatial normalization to Montreal Neurological Institute (MNI) space using unified seg-
mentation method, and spatial smoothing with a 6 mm full-width at half-maximum Gaussian
kernel. A high-pass filter cut-off was set as 128 s.
At the first level analysis, we modeled each block (five trials from the same condition, duration:
30 s) as an event with a general linear model in which the time series was convolved with the canon-
ical hemodynamic response function (HRF) with its temporal and dispersion derivatives (Brouwer
& Heeger, 2013; Friston et al., 2011; Ward et al., 2008). The derivatives were included because the
Sun et al. 201

HRF of an individual voxel may have differed from the canonical HRF. The variance of the esti-
mated response amplitudes across runs was smaller with the derivative included than without it
(Brouwer & Heeger, 2013). Nuisance regressors consisted of six head motion parameters and a con-
stant regressor for each run.

Contrasts. First, we investigated the effect of the tracking task on brain activities by comparing the
pooled data from all experimental blocks with an implicit baseline. The implicit baseline refers to
the brain activities during the unmodeled time when no stimuli were presented, including the inter-
block interval and rest period.
Second, to investigate the effects of hue distance and category separately, we compared the
Large condition versus the Small condition (across within- and cross-category) and the Large
Within condition versus the Large Cross condition.
Linear contrasts of coefficients for each participant were entered into second level
random-effects analyses. Group statistics including the canonical HRF and its temporal and disper-
sion derivatives were computed for each contrast, resulting in 15 pairs of contrasts (canonical: task
> baseline, Large > Small, Small >Large, LW > LC, LC > LW; temporal derivative: task > baseline,
Large > Small, Small >Large, LW > LC, LC > LW; dispersion derivative: task > baseline, Large >
Small, Small >Large, LW > LC, LC > LW). The cluster-wise threshold was set to control the
family-wise error (FWE) rate at p < .05. A primary voxel-level threshold was set as p < .001 to
define clusters. Cluster size > = 20.

Correlation Analysis. We also conducted brain-behavior correlation analyses to investigate whether

the brain activities in cross-category conditions for participants were dependent on their subject-
ive/perceived categorical distance for the color stimuli. The rationale for this analysis was
twofold. First, there are considerable individual differences in color categorization in the literature
(e.g., Brown et al., 2016; Gibson et al., 2017) and our study. Thus, the comparison between cross-
and within-category conditions based on the group-level category boundary might have masked
the actual categorical effects. Second, it is a major concern that the perceptual color differences
are not fully controlled when testing the effects of color categories (for review, see Witzel, 2019).
The correlation between brain activities and subjective/perceived categorical distance (if there
was one) would demonstrate that the categorical effects were not due to low-level color differences.
In this analysis, we performed a subject-level simple linear regression analysis using linear con-
trasts of coefficients from first-level analysis, reflecting the effect of the Large Cross conditions
versus baseline as the dependent variable and perceived categorical distance as a regressor. The
regression analysis was conducted at the whole-brain level, separately for the canonical HRF, its tem-
poral, and dispersion derivatives. The cluster-wise threshold was set to control the FWE rate at p <
.05. A primary voxel-level threshold was set as p < .001 to define clusters. Cluster size > = 20.

Regions of Interest Analysis. Both functional and structural Regions of Interest (ROIs) are defined.
First, we found that the bilateral middle temporal complex (MT + /V5, peaking at [−45, −69, 6]
and [48, −66, 3]) showed larger BOLD amplitudes during tracking (Figure 1C, Table 1). As our
study did not conduct a motion localizer task, we compared the locations of our MT + with
those reported in the literature (Supplemental Material, Table S1). It appears that our MT +
largely overlaps with the previously reported MT + . The MT + is a motion-sensitive area whose
involvement has been reported in previous MOT studies (Chakraborty et al., 2021; Culham
et al., 1998; Howe et al., 2009; Merkel et al., 2015; Nummenmaa et al., 2017). Accordingly, we
defined two separate functional ROIs (left MT + and right MT + ) as two spheres with a radius
of 6 mm centered at the peak voxels in the significant clusters. Note that the ROIs defined were
not biased, concerning the tests of interest (e.g., whether there are significant differences
202 Perception 52(3)

Table 1. Coordinates of activation peaks: task > baseline.

MNI coordinates

Brain region BA Cluster size x y z T

Task > Baseline (HRF)

L MT + 37 220 −45 −69 6 11.18
19 −46 −74 1 6.09
R MT + 37 258 48 −66 3 11.01
19 45 −77 1 4.88

between the Large Within condition and the Large Cross condition). In addition, we defined struc-
tural ROIs based on a previous study (Purmann & Pollmann, 2015) that reported the MNI coordi-
nates of peak voxels of four color-selective areas. According to this study, four ROIs (Figure 1D),
namely, the left V4, right V4, left V4a, and right V4a, were defined as spheres with a radius of 6 mm
[peak voxel: (−30, −73, −11), (31, −70, −11), (−29, −55, −13), and (29, −47, −17), respectively].
For each ROI, we conducted two types of analyses: univariate analyses and representational simi-
larity analyses (RSA; Kriegeskorte et al., 2008). For univariate analyses, the percent signal change in
each condition was obtained for each subject using the MarsBaR toolbox (Brett et al., 2002). The
differences in the percent signal change between the Large conditions and the Small conditions
(across within- and cross-category) and between the Large Within condition and the Large Cross
condition were analyzed using paired t-tests for each ROI. For RSA, the t-value images of four con-
ditions, reflecting the effect of the condition on magnitude of the BOLD response, were calculated to
capture the activation patterns, which were generated from unsmoothed data. For each region, the
activation patterns within the ROI were extracted to calculate the neural representational dissimilar-
ity matrix (RDM) using the CoSMoMVPA toolbox (Oosterhof et al., 2016). A 4 × 4 neural RDM was
obtained for each region and each participant, where the dissimilarity measure was computed by one
minus Pearson correlation (range: 0–2). For the RSA analysis of categorical representation, we inves-
tigated the similarity in patterns of activity between conditions with the same category relationship
(LW-SW, LC-SC), with the similarity in patterns of activity between conditions with different cat-
egory relationships (LW-SC, LC-SW). Specifically, we investigated whether the average similarity
between LW and SW and between LC and SW was larger than the average similarity between LW
and SC and between LC and SW. The size of hue distance was balanced for this comparison.
Similarly, for the RSA analysis of the representation of hue distance, we investigated the similarity
in patterns of activity between conditions with same hue distance (LW-LC, SW-SC), with the simi-
larity in patterns of activity between conditions with different hue distances (LW-SC, LC-SW).
Specifically, we investigated whether the average similarity between LW and LC and between
SW and SC was larger than the average similarity between LW and SC and between LC and SW.
The category relationship was balanced for this comparison.

Results
Behavioral Results
The mean tracking accuracy (ACC; the proportion of responding correctly to whether the probe was
presented or not) and reaction time (RT; only trials with correct response included) were analyzed.
As shown in Figure 2A, the tracking ACC in the Small conditions was significantly larger than that
in the Large condition (within- and cross-category trials were averaged first for each participant; t
Sun et al. 203

(16) = 2.20, p = .043, Cohen’s d = 0.54). Large trials showed no significant differences in tracking
ACC between the Within condition and the Cross condition (Large Within vs. Large Cross: t (16) =
0.17, p = .871, Cohen’s d = 0.04). For RTs (Figure 2A), there was no significant difference between
the Large condition and the Small condition (t (16) = 1.27, p = .222, Cohen’s d = 0.31). By contrast,
the RT was significantly shorter in the Large Within condition than the Large Cross Condition (t
(16) = 3.15, p = .006, Cohen’s d = 0.76). These results suggested that both the hue distance and cat-
egorical relationship of the targets modulate the tracking process. The targets were easily grouped
together to ease tracking when the hue distance of the targets is smaller and when they are from the
same category, consistent with previous findings (Erlikhman et al., 2013; Howe & Holcombe,
2012; Lupyan, 2008; Sun et al., 2020). As in Sun et al. (2020), the CP effects in tracking are man-
ifested in within-category facilitation instead of between-category facilitation in almost all other
studies, for smaller distinctiveness within the target set facilitates tracking.

fMRI Results
Contrasts
The effect of hue distance was found in the medial prefrontal areas (Small > Large, dispersion
derivative), that is, shorter duration of hemodynamic responses in the Small condition than the
Large condition (Table 2, Figure 2B). The differences were mainly observed in medial regions
of the bilateral Superior Frontal Gyrus (SFGmed). The SFGmed was previously shown to
govern high-level cognitive functions (Ridderinkhof et al., 2004; Rushworth et al., 2004).
Studies of attentive tracking have shown the involvement of this area (e.g., Nummenmaa et al.,

Figure 2. Behavioral and fMRI results. (A) Mean task performance (ACC and RT) in the behavioral task. (B)
Brain regions showing hue effects (Small > Large) in the dispersion derivative. Error bars show ± SEM. (C)
Brain regions showing category effects (Large Cross > Large Within) in the time derivative. (D) Brain regions
showing significant correlation between the categorical distance and the time derivative in the Large Cross
condition. Warm colors indicate positive correlation, and cool colors indicate negative correlation. The
scatter plot shows the voxel-averaged t-values against categorical distances in the SFGmed (red) and MTG
(blue).
204 Perception 52(3)

2017). The differences between the Large and Small conditions in these frontal areas suggested that
the hue distance among moving objects could mediate high-level processes during tracking.
In addition, the categorical effects were revealed in the right lateral prefrontal areas (i.e., MFG)
and the posterior inferior parietal lobule (IPL). In these areas, earlier hemodynamic responses were
observed in the Large Cross condition compared to the Large Within Condition (LC > LW, time
derivative; Table 2, Figure 2C).

Correlation
It was found that, for the Large Cross condition, the categorical distance was negatively correlated
with the dispersion/duration of hemodynamic responses in the left middle temporal gyrus (MTG)
and also positively correlated with the HRF dispersion in the right SFGmed (Table 3, Figure 2D).
The MTG identified here is located anterior to the MT + ROIs and lateralized to the left hemisphere.
The left MTG likely supports language processing of the color stimuli, and there are individual dif-
ferences in verbal/categorical encoding for one same color pair.

ROI Analyses
Univariate Analyses. The ROI-based analyses on the percent signal change revealed significant categor-
ical effects in left MT + and right V4 (Figure 3A). The percent signal change in the Large Within
condition was significantly larger than that in the Large Cross condition in the left MT + (t (16) =
2.49, p = .024, Cohen’s d = 0.60) and right V4 (t (16) = 2.13, p = .049, Cohen’s d = 0.52). There
were no significant differences between Large Cross and Large Within in any other ROIs (ps >
.05). In addition, there were no significant differences between the Large conditions and the Small con-
ditions (within and cross trials are averaged for each participant) in any ROIs (ps > .05).

Representational Similarity Analyses. Figure 3B shows the group-level neural RDMs for six ROIs.
Overall, these ROIs showed similar patterns in neural RDMs. For example, the similarity in activity
patterns between Large Within and Large Cross is the greatest for all ROIs. Using paired t-tests, we

Table 2. Coordinates of activation peaks: Small > Large, Large Cross > Large Within.

MNI coordinates

Brain region BA Cluster size x y z T

Small > Large (dispersion derivative)

L superior medial gyrus 8 542 −3 39 48 4.99
9 −3 51 33 4.67
L superior frontal gyrus 9 −9 51 39 4.6
Large Cross > Large Within (time derivative)
R middle frontal gyrus 48 89 39 33 21 4.21
46 39 48 6 3.70
45 45 42 18 3.30
R inferior frontal gyrus (triangular part) 48 39 18 21 3.43
R inferior parietal lobule 39 138 39 −57 45 4.38
R superior parietal lobule 7 39 −60 60 3.51
R angular 39 39 −57 48 4.13
R thalamus 109 9 −15 0 4.14
12 −12 15 4.11
Sun et al. 205

Table 3. Coordinates of activation peaks: correlation between brain activation and categorical distance.

MNI coordinates

Brain region Brodmann area Cluster size x y z T

Positive correlation (dispersion derivative)

R superior medial gyrus 9 87 6 45 48 8.74
Negative correlation (dispersion derivative)
L middle temporal gyrus 37 66 −60 −57 3 6.14

Figure 3. ROI results. (A) Mean percent signal changes in the six ROIs. Error bars show ± SEM. (B) Color
map showing the neural distance in activation patterns between every two conditions in the six ROIs at the
group level.

found that the similarity in activity patterns between conditions with same category relationship
(LW-SW, LC-SC) was larger than the similarity between conditions with different category rela-
tionships (LW-SC, LC-SW) in bilateral MT + (left MT + : t (16) = 4.47, p < .001, Cohen’s d =
1.09; right MT + : t (16) = 3.56, p = .003, Cohen’s d = 0.86) and left V4 (t (16) = 3.01, p = .008,
Cohen’s d = 0.73). Other ROIs showed no such effects. The results suggested that the categorical
relationship of the color stimuli is encoded in bilateral MT + and left V4 areas.
In addition, the similarity in activity patterns between conditions with same hue distance
(LW-LC, SW-SC) was larger than the similarity between conditions with different hue distances
(LW-SC, LC-SW) in bilateral MT + (left MT + : t (16) = 4.47, p < .001, Cohen’s d = 1.09; right
MT + : t (16) = 3.56, p = .003, Cohen’s d = 0.86), bilateral V4 (left V4: t (16) = 5.53, p < .001,
Cohen’s d = 1.34; right V4: t (16) = 4.12, p < .001, Cohen’s d = 1.00), and bilateral V4a (left
V4a: t (16) = 3.08, p = .007, Cohen’s d = 0.75; right V4a: t (16) = 3.90, p < .001, Cohen’s d =
0.95). These findings revealed that bilateral MT + , bilateral V4, and bilateral V4a carry the infor-
mation of hue distance between the moving color stimuli.

Discussion
The current study investigated the neural mechanism behind the categorical effect of color in MOT.
We found that categorical effects of color in MOT were associated with activities in a broad range
of brain regions, including both the ventral (V4, MTG) and dorsal pathways (MT + , IPL) of feature
processing, as well as frontal regions (MFG, SFGmed). We proposed that these regions are
206 Perception 52(3)

hierarchically organized and responsible for distinct functions. The color-selective V4 encodes
color categories, making cross-category colors more discriminable than within-category colors
(Brouwer & Heeger, 2013; Sun et al., 2021). Meanwhile, the language or semantic regions
encode the verbal information of the colors. Both visual and nonvisual codes of color categories
then modulate the activities of motion-sensitive MT + areas and frontal areas responsible for atten-
tional processes.

Visual and Nonvisual Categorical Color Encoding. Whether categorical effects occur at the perceptual
level has been the focus of argument among CP studies (for review, see Regier & Kay, 2009;
Siuda-Krzywicka et al., 2019; Witzel, 2019). Using RSA, we found that color stimuli with the
same categorical relationship (e.g., within-category vs. within-category) exhibited more similarity
in activation patterns than those with different categorical relationships (within-category vs. cross-
category) in left V4. Notably, in both bilateral V4 and V4a, color stimuli with the same hue distance
exhibited more similarity in activation patterns than those with different hue distances. These find-
ings suggested that the defined color-selective ROIs are sensitive to perceptual color differences and
that the high-order visual cortex carries the information of color categories, consistent with some
recent findings (Brouwer & Heeger, 2013; Siok et al., 2009; Sun et al., 2021). Compared to the
hypothesis that the perceptual color space is inbuilt categorically in the human brain, recent
fMRI studies point to the more plausible possibility that high-level cognition (e.g., attention)
exerts top-down modulation on color perception and that categorical color perception is task-
modulated (Brouwer & Heeger, 2013; Siok et al., 2009; Sun et al., 2021). This top-down view
may explain our findings here because MOT is an attention-consuming task, revealed by the
involvement of medial frontal areas (SFGmed). However, we cannot exclude the possibility that
the perceptual color space is inbuilt categorically in the human brain. In addition, the signal
change was greater in the Large Cross condition than in the Large Within condition in right V4,
which is in line with the findings of Siok et al. (2009).
In addition to the categorical representation of color in the visual cortex, categorical encoding of
color was also associated with brain activities in the left MTG (anterior to MT + ), right IPL, right
MFG. At the whole-brain level, tracking cross-category targets resulted in earlier hemodynamic
responses in the right IPL and MFG, compared to within-category targets. Moreover, the categor-
ical distance was negatively correlated with the dispersion of hemodynamic responses in the left
MTG. The involvement of these areas is unique for CP effects and not shared by hue effects.
All these areas have been previously reported to be associated with language or semantic processing
(for review, see Jackson, 2021). These results may suggest that language- and semantic-processing
areas are directly involved in encoding color categories in the tracking task, even though the task
requires no explicit color judgment. The involvement of language-related processes in categorical
effects is consistent with some previous studies (Bird et al., 2014; Siok et al., 2009; Tan et al.,
2008). Both Tan et al. (2008) and Siok et al. (2009) found that language processing areas were
engaged in perceptual discrimination of colored stimuli. The participation of the left MTG in our
study likely supports verbal labeling of the moving colors and that individual differences were
encoded in the brain activity of the left MTG.
In contrast to the MTG, the comparison between Large Cross versus Large Within revealed the
involvement of MFG and IPL in the right instead of the left hemisphere. This result may suggest
that part of the CP effects in tracking is language-independent. Although the CP effects initially
serve as a bridge between language and perception, recent evidence has been found that CP can
be language-independent (e.g., Siuda-Krzywicka et al., 2019) and that it occurs in either visual
field (Siok et al., 2009; Witzel & Gegenfurtner, 2011, 2015, 2016) or is even lateralized in the
left visual field (Suegami et al., 2014). Bird et al. (2014) also observed MFG showing greater
Sun et al. 207

activity for cross-category colors relative to within-category colors in passive viewing, supporting a
domain-general categorization underlying categorical effects. Different from our study, the activa-
tion in MFG in their study was bilateral. Studies of categorization in other domains have suggested
right lateralization of the MFG in categorical processing (e.g., Sachs et al., 2008). Also, the IPL has
been related to the memory system (for review, see Igelström & Graziano, 2017), and it has often,
but not always, been left-lateralized (e.g., Summerfield et al., 2009). The engagement of the right
MFG and IPL may support domain-general categorical representation at the conceptual level.
Based on the left-hemisphere-lateralized activation in the MTG and right- hemisphere-lateralized
activation in the MFG and IPL, our study points to both language-dependent CP and
language-independent CP in tracking. It is in line with the previous behavioral study, which
found that verbal interference disrupts only part of the CP effects in tracking (Sun & Zhang, 2022).

The Effect of Color Categories on Tracking. Our study also aimed to uncover how color categories
influence the tracking process. We suggested that the MT + and medial prefrontal areas are
likely responsible for the modulation of both color categories and hue distance on tracking. The
ROI analyses revealed that the signal changes in left MT + were greater in the Large Within con-
dition than in the Large Cross condition. For the RSA analyses, both the categorical relationship
and the hue distance influence the activation patterns in the MT + regions. MT + is a motion-
selective area whose involvement has been repeatedly reported in MOT studies (Chakraborty
et al., 2021; Culham et al., 1998, 2001; Howe et al., 2009; Merkel et al., 2015; Nummenmaa
et al., 2017). MT + is associated with many different types of motion percepts. For example,
MT + is sensitive to the proportion of items that are moving coherently (Braddick et al., 2001;
Rees et al., 2000). Local and global motion (Strong et al., 2017) and imagined motion (Goebel
et al., 1998) can all invoke the activities of MT + . In addition to motion processing, MT + is
also engaged in the processing of motion-defined form (Strong et al., 2017) and object shape
(Kourtzi et al., 2002). It may result from the overlapping in cortical regions between MT + and
LO, which is responsible for object recognition. In our study, the six moving stimuli have two
surface colors, and the featural difference or similarity could easily affect the motion percepts.
When these two colors are distinct from each other, two sets of moving stimuli would automatically
be formed based on feature-based grouping, and the motion pattern becomes more local (i.e., two
independently moving groups). When these two colors are similar to each other, all the moving
stimuli can be grouped into a single virtual object representation (Yantis, 1992), and the motion
pattern becomes more global. We suggested that both hue distance and categorial relationship influ-
ence the moving pattern in this way, and that is why we observed that these two kinds of informa-
tion are encoded in the activation patterns of MT + . More accurately, the MT + does not encode the
color features per se but encodes the motion patterns, which are modulated by color features.
In addition, the Large Within condition showed increased signal changes than the Large Cross
condition. We suggested two possibilities behind this finding. One of them is that the difference in
signal changes might be an attention effect. There is evidence showing the MT + response is sen-
sitive to attention (e.g., Berman & Colby, 2002; Huk et al., 2001). In our study, the difference in
categorical relationship between conditions may encourage the participants to attend more to the
moving colors in the Large Within condition because of the conflicting perceptual and categorical
codes (i.e., difference hues, same category). Consequently, the Large Within condition exhibited
increased signal changes than the Large Cross condition. Alternatively, the difference in BOLD
signal changes is a genuine category effect rather than an attention effect. In the Large Within con-
dition, the large perceptual differences automatically segment the stimuli into two groups, while the
shared category membership can disrupt this process. This conflicting information could place dif-
ficulty in perceiving the motion patterns and consequently increase activities in MT + .
208 Perception 52(3)

The engagement of the medial SFG was observed in both CP and hue effects. SFGmed governs
high-level cognitive functions, such as attention shift (Nagahama et al., 1999), action selection
(Rushworth et al., 2004), and conflict monitoring (Ridderinkhof et al., 2004). Several recent studies
of attentive tracking have shown the involvement of this area (Nummenmaa et al., 2017; Wang
et al., 2018). Wang et al. (2018) found decreased activation in left SFGmed in multiple grouping con-
ditions where the targets can be grouped together compared to no-grouping conditions. The activity of
SFGmed seems sensitive to the tracking demand (Wang et al., 2018). The activity of SFGmed (the
HRF dispersion) is sensitive to both categorical differences and hue distance in our study, which
may suggest that the brain response in this area is associated with behavioral tracking performance.
There are several limitations of our study. One limitation is that the involvement of several brain
areas was observed in the latency and dispersion of the BOLD responses rather than in the magni-
tude. It might be worrying that we directly related the brain regions with differences in the temporal
characteristics of BOLD responses with those in BOLD amplitude from previous studies. However,
we believe the comparison is meaningful for two reasons. First, recent evidence suggests that the
temporal structure of BOLD signals is correlated with the amplitude (Grinband et al., 2008;
Thompson et al., 2014). For example, Thompson et al. (2014) found that larger neural responses
produce BOLD signals that begin earlier in time. The earlier onset of BOLD signals could mean
earlier peak latency or longer duration. It is reasonable to deduce that the differences in temporal
characteristics and amplitudes of BOLD responses may have similar neural sources. Our study ana-
lyzed the fMRI data with SPM (Statistical Parametric Mapping) which applies automatic orthogo-
nalization when derivatives of regressors are added to a model. Therefore, this method does not
allow us to reveal the correlation between the regressors. However, it does not necessarily mean
that the brain regions showing differences in temporal characteristics must be isolated from the
brain regions showing differences in BOLD amplitudes. There are also previous studies that dir-
ectly compared their results in temporal characteristics with the results in the amplitude (i.e.,
Henson et al., 2002; Ward et al., 2008). Second, the brain areas showing differences in latency
or duration of BOLD responses in our study have also been reported in previous studies.
Previous evidence suggested that the language-related regions produce larger, delayed (in peak
latency), and more dispersed BOLD responses to nonwords (Henson et al., 2002) or rotated
words (Bellgowan et al., 2003), because of the increased difficulty in searching for a lexical
representation. In our study, the language/semantic processing areas (e.g., left MTG) showed
earlier BOLD responses for the Large Cross stimuli relative to the Large Within stimuli. The
delayed BOLD response for the Large Within stimuli in our study may suggest that labeling two
distinct colors with the same lexical term is more challenging than labeling them with two different
lexical terms. In addition, it was suggested that the duration of BOLD responses in the medial
frontal cortex is associated with the duration of the stimuli (Waugh et al., 2010; 2014). The differ-
ences in the duration of neural responses across different conditions may result from the varying
durations of attention maintenance during tracking.
Another limitation is that our study used a limited number of color samples. In the within-category
conditions, only green color pairs are used, resulting in a greater proportion of green colors than blue
colors overall. In the previous CP studies using MOT (Sun et al., 2020, 2018), the amount of green and
blue colors is the same. Despite the differences in color sampling, our behavioral results are consistent
with the findings from these studies. We cautiously concluded that the current color sampling is accept-
able, but a more balanced sampling would be more appropriate.

Acknowledgement
This work was supported by the General Program of National Natural Science Foundation of China
(31271083) (to XZ) and the Key Program of National Natural Science Foundation of China (61632014) (to
XZ).
Sun et al. 209

Author Contribution(s)
Mengdan Sun: Conceptualization; Data curation; Formal analysis; Investigation; Methodology;
Visualization; Writing – original draft; Writing – review & editing.
Xiaoyang Xin: Data curation; Formal analysis.
Haojiang Ying: Methodology; Writing – review & editing.
Luming Hu: Investigation; Writing – review & editing.
Xuemin Zhang: Conceptualization; Formal analysis; Resources; Supervision; Writing – review & editing.

Declaration of Conflicting Interests

The authors declared no potential conflicts of interest with respect to the research, authorship, and/or publica-
tion of this article.

Funding
The authors disclosed receipt of the following financial support for the research, authorship, and/or publication
of this article: This work was supported by the Scientific and Technological Innovation 2030 (No.
2021ZD0204300), National Natural Science Foundation of China (grant numbers 31271083, 61632014).

ORCID iDs
Mengdan Sun https://orcid.org/0000-0002-1293-3006
Haojiang Ying https://orcid.org/0000-0002-6371-6650
Xuemin Zhang https://orcid.org/0000-0002-5357-6666

Supplemental Material
Supplemental material for this article is available online.

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