Journal of Vision (2019) 19(12):22, 1–14
Category selectivity for animals and man-made objects:
Beyond low- and mid-level visual features
Chenxi He
Olivia S. Cheung
Distinct concepts, such as animate and inanimate
entities, comprise vast and systematic differences in
visual features. While observers search for animals faster
than man-made objects, it remains unclear to what
extent visual or conceptual information contributes to
such differences in visual search performance. Previous
studies demonstrated that visual features are likely
sufficient for distinguishing animals from man-made
objects. Across four experiments, we examined whether
low- or mid-level visual features solely contribute to the
search advantage for animals by using images of
comparable visual shape and gist statistics across the
categories. Participants searched for either animal or
man-made object targets on a multiple-item display with
fruit/vegetable distractors. We consistently observed
faster search performance for animal than man-made
object targets. Such advantage for animals was unlikely
affected by differences in low- or mid-level visual
properties or whether observers were either explicitly
told about the specific targets or not explicitly told to
search for either animals or man-made objects. Instead,
the efficiency in categorizing animals over man-made
objects appeared to contribute to the search advantage.
We suggest that apart from low- or mid-level visual
differences among categories, higher-order processes,
such as categorization via interpreting visual inputs and
mapping them onto distinct concepts, may be critical in
shaping category-selective effects.
Introduction
We maintain stable representations of the world via
interactions of incoming sensory input and existing
conceptual knowledge about the world. Distinct con-
cepts, such as animate and inanimate entities, often
comprise vast and systematic differences in visual
features. As the brain transforms percepts into concepts,
to what extent do the mental representations that guide
our behavior in searching for relevant items in an
Citation: He, C., & Cheung, O. S. (2019). Category selectivity for animals and man-made objects: Beyond low- and mid-level
visual features. Journal of Vision, 19(12):22, 1–14, https://doi.org/10.1167/19.12.22.
https://doi.org/1 0. 11 67 /1 9 .1 2. 22 Received June 13, 2019; published October 24, 2019
This work isonlicensed
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1
Department of Psychology, Division of Science,
New York University Abu Dhabi, United Arab Emirates
Department of Psychology, Division of Science,
New York University Abu Dhabi, United Arab Emirates $
environment contain visual or conceptual information?
While several recent studies have shown that the effects
of visual properties may account for category effects
such as animacy (e.g., Long, Störmer, & Alveraz, 2017;
Zachariou, Del Giacco, Ungerleider, & Yue, 2018; see
also Rice, Watson, Hartley, & Andrews, 2014), here we
examine whether higher-level processes such as catego-
rization may also influence visual search performance
when low- and mid-level visual differences that have
been previously shown to sufficiently distinguish among
animals and man-made objects are minimized.
One animate–inanimate distinction in visual per-
ception is that there is an advantage for the processing
of animals compared with man-made objects. For
instance, observers are faster at detecting animals,
compared with man-made objects or other inanimate
items such as plants, in complex natural scenes (New,
Cosmides, & Tooby, 2007; Wang, Tsuchiya, New,
Hurlemann, & Adolphs, 2015). Even when observers
are not actively searching for animals, the presence of
animals in a scene also appears to capture attention.
For instance, change detection for inanimate targets
(e.g., a leaf) was slowed when there were animal
distractors in the display, compared with when there
were no animal distractors (Altman, Khislavsky,
Coverdale, & Gilger, 2016). The attentional advantage
for animals has also been shown in other tasks
examining visual search efficiency (Jackson & Calvillo,
2013; Lipp, Derakshan, Waters, & Logies, 2004; but see
Levin, Takarae, Miner, & Keil, 2001), resistance of the
inattentional blindness (Calvillo & Hawkins, 2016;
Calvillo & Jackson, 2014) and attentional blink
(Guerrero & Calvillo, 2016), using behavioral or eye
tracking measures (Yang et al., 2012).
Despite the wealth of evidence of the attentional
advantage for animals compared with man-made
objects, it remains unclear whether visual or conceptual
aspects of the differences between animals and man-
made objects may contribute to the advantage. For
ISSN 1534-7362 Copyright 2019 The Authors
 Journal of Vision (2019) 19(12):22, 1–14
complex natural scenes, although systematic differences
in low-level visual properties, such as power spectrum,
luminance, and contrast, have been found between
images with or without animals in the scenes (Torralba
& Oliva, 2003; Wichmann, Drewes, Karl, & Gegen-
furtner, 2010), such differences cannot account for
human observers’ rapid detection of animals in these
images (Wichmann et al., 2010). Nonetheless, inde-
pendent of any scene details, it is possible that visual
features of the items alone are sufficient for human
observers to distinguish between animals and man-
made objects (e.g., Levin et al., 2001; LoBue, 2014).
Specifically, animals often have curvilinear shapes
whereas man-made objects, especially tools, tend to be
rectilinear or elongated in shape, which affords
graspability (Almeida et al., 2014). Curvilinear and
rectilinear visual features alone may be sufficient to
support categorization between animals and man-made
objects even for scrambled, texture-form images with or
without recognizable global shape information (Long
et al., 2017; Zachariou et al., 2018). Such mid-level
visual features facilitate visual search performance if a
target (e.g., animal) is from a different category than
the distractors (e.g., man-made objects; Long et al.,
2017). Animals and man-made objects also differ in
visual similarity among category members (Gerlach,
Law, & Paulson, 2004), as animals tend to be more
visually similar to each other (e.g., many animals have
four legs and one head, etc.) than man-made objects
(e.g., different tools may have few overlapping visual
features; Humphreys, Riddoch, & Quinlan, 1988).
Moreover, animals may also be more visually complex
compared with man-made objects (e.g., a lion vs. a
hammer; Gerlach, 2007; Moore & Price, 1999; Snod-
grass & Vanderwart, 1980; Tyler et al., 2003). Taken
together, it is possible that vast differences in the visual
features between animals and man-made objects may
drive the differential performance in visual search.
If the advantage for animals in visual search remains
with images of comparable visual features between the
categories, it would suggest that higher-level processes
such as categorization, which involves semantic pro-
cessing or extracting meanings from any available
visual features, may play an additional role. Here we
tested this hypothesis by minimizing visual differences
among the categories. Our findings would extend from
previous studies that found search advantage for
animals to further understand the nature of represen-
tations among animals and man-made objects that are
critical for performance.
The current study used two approaches in attempt to
minimize visual differences between animals and man-
made objects. First, we selected images that were either
round or elongated in the outline shapes. Second, we
measured gist statistics of the images and used only
images with comparable gist statistics between the
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He & Cheung 2
categories. Gist statistics describe the holistic shape and
structure of an image by sampling spatial frequency
and orientation information of image segments (Oliva
& Torralba, 2001, 2006), which capture more complete
and sophisticated visual shape properties compared
with other low-level visual measures such as luminance,
contrast, or pixel-wise measures. Gist statistics also
provide information about the amount of visual details
in the images (Oliva & Torralba, 2006). More
importantly, gist statistics have been shown to support
behavioral performance in visual categorization, and to
predict neural responses in the occipitotemporal cortex
for various visual object and scene categories (e.g.,
Andrews, Watson, Rice, & Hartley, 2015; Bar, 2003;
Loschky & Larson, 2010; Oliva & Torralba, 2001,
2007; Rice et al., 2014; Watson, Hartley, & Andrews,
2014). Here, we used images of comparable visual
shape and gist statistics across different categories to
examine whether the search advantage for animals
would remain.
In Experiments 1A, 1B, and 2A, observers were
asked to search for either an animal or a man-made
object target on a multiple-item display with varied set
sizes, with fruits/vegetables as additional fillers. In
Experiment 2B, observers were told to search for any
non-fruits/vegetable items, which could either be
animals or man-made objects. Note that in addition to
comparable visual shape and gist statistics in the
images we used across the three categories, fruits/
vegetables were used as additional fillers as previous
studies showed comparable semantic distance of fruits/
vegetables to either animals or man-made objects
(Bracci & Op de Beeck, 2016; Carota, Kriegeskorte,
Nili, & Pulvermüller, 2017).
Our main focus was the effect of animacy on the
search for a target, in other words, whether the search
for animals is faster than the search for man-made
objects among items with comparable visual shape and
gist statistics. Apart from the main focus of the effect of
target category, in Experiments 1A and 1B we also
examined whether task-irrelevant animals or man-
made objects may attract attention away from the
target, revealing a stimulus-driven effect (Langton,
Law, Burton, & Schweinberger, 2008; Theeuwes, 1991,
1992; Yantis, 1993). Therefore, in half of the trials, an
item from the non-target category (e.g., a man-made
object when searching for an animal) also appeared as a
distractor among the fruit/vegetable distractors. Addi-
tionally, while we do not expect animacy to be a basic
feature that guides visual search (Wolfe & Horowitz,
2004; but see Levin et al., 2001), as indicated by a pop-
out effect for animal search with the search slope less
than 10 ms/item with respect to the set size increase
(Duncan & Humphreys, 1989; Theeuwes, 1993; Treis-
man & Gelade, 1980; but see Buetti, Cronin, Madison,
Wang, & Lleras, 2016), we examined the search
 Journal of Vision (2019) 19(12):22, 1–14 He & Cheung
performance for animal and man-made object targets
across set sizes of 3, 6, and 9 in all experiments.
To anticipate the results, we found in Experiment 1A
that the search was faster for animal than man-made
object targets across set sizes. In Experiment 1B, we
replicated the results in Experiment 1A while further
ruling out the influences of low-level visual factors
including luminance, contrast, and power spectrum, as
these factors could affect selective attention (Moraglia,
1989; Parkhurst, Law, & Niebur, 2002; Sagi, 1988;
Smith, 1962; Theeuwes, 1995; Wolfe & Horowitz, 2004,
2017). Experiments 2A and 2B further demonstrated
that categorization is more efficient for animals than
man-made objects, either when participants were
shown the names of all target items prior to the search
task to minimize the range of possible targets for both
categories, or when participants were asked to detect
any items that were not fruits/vegetables.
Experiments 1A and 1B
Method
Participants
Fifty-six undergraduate students aged between 18
and 23 of years (mean ¼ 19.8, SD ¼ 1.3, 34 women and
22 men) at New York University Abu Dhabi partici-
pated for either course credits or subsistence allow-
ances; 28 participants completed Experiment 1A and 28
participants completed Experiment 1B. All participants
had normal or corrected-to-normal vision. All exper-
iments were approved by New York University Abu
Dhabi Institutional Review Board.
Stimuli
Figure 1A illustrates examples of the stimuli in
Experiment 1A. A total of 52 items including 12
animals, 12 man-made objects, and 28 fruits/vegetables
were used. Each item had 16 exemplars. All images
were in grayscale. An exemplar of each item is shown in
the Appendix, Figure A1. The animals and man-made
objects were either target or distractor categories for
each of the two participant groups. The fruits/
vegetables were filler distractors. Half of the items from
the three categories had round shapes (e.g., turtle,
steering wheel, apple), while the other half had
elongated shapes (e.g., wall lizard, butter knife,
cucumber). All items had neutral valence, so any
potential emotional effect on visual search should be
minimized (e.g., LoBue, 2014; Ohman, Flykt, &
Esteves, 2001).
Gist statistics were measured on spatial frequency
and orientation information of all segments for each
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3
Figure 1. Sample images of animals, man-made objects, and
fruits/vegetables used in Experiments 1A (A) and 1B (B). In all
three categories, the overall shape of half of the items was
elongated, whereas the overall shape of the remaining items
was round.
image (Oliva & Torralba, 2001). Specifically, a series of
Gabor filters across eight orientations and four spatial
frequencies were applied to each image to generate 32
filtered images. Each filtered image was then segmented
into a 4 3 4 grid within which the values were averaged
per cell, resulting in 16 values. The final gist statistics
for each image were made up by a vector of 512 (32 3
16) values (e.g., Oliva & Torralba, 2001; Rice et al.,
2014). To compare gist statistics across shapes and
categories, the values were first averaged across all
exemplars for each item. Dissimilarity indicated by
squared Euclidean distance between gist statistics of
each pair of items both within and across shapes and
categories were then calculated (Figure 2). We com-
pared the dissimilarity between round versus elongated
shapes separately for each category, and between each
pair of categories separately for each shape. We found
that the gist statistics were significantly different
between elongated and round shapes in each category,
but they were comparable across categories for either
shape. Within-shape dissimilarity (e.g., a turtle and a
squirrel) was significantly lower than cross-shape
dissimilarity (e.g., a turtle and a wall lizard) for all
categories: animals, t(64) ¼ 5.9, p , 0.0001; man-
made objects, t(64) ¼ 12.7, p , 0.0001; fruits/
vegetables, t(376) ¼ 11.8, p , 0.0001. Conversely,
there was no statistical difference between within-
category dissimilarity (e.g., a turtle and a squirrel) and
cross-category dissimilarity (e.g., a turtle and a steering
wheel) for animals versus man-made objects: elongated,
 Journal of Vision (2019) 19(12):22, 1–14 He & Cheung 4

Figure 2. Pair-wise dissimilarity (squared Euclidean distance) of gist statistics showed significant differences between elongated and
round shapes in each category, but no significant differences across animals, man-made objects, and fruits/vegetables for either
shape.

t(64) ¼0.2, p ¼ 0.82; round, t(64) ¼0.02, p ¼ 0.99, for
animals versus fruits/vegetables: elongated, t(188) ¼
0.4, p ¼ 0.71; round, t(188) ¼ 0.6, p ¼ 0.52, and for man-
made objects versus fruits/vegetables: elongated, t(188)
¼ 0.6, p ¼ 0.58; round, t(188) ¼ 0.6, p ¼ 0.58. These
results indicated that for either elongated or round
shape, there was no systematic difference across
different categories in visual properties quantified by
gist statistics.
Low-level visual features, such as luminance, con-
trast and power spectrum could affect selective
attention (Moraglia, 1989; Parkhurst et al., 2002; Sagi,
1988; Smith, 1962; Theeuwes, 1995; Wolfe & Horowitz,
2004). For the images used in Experiment 1A, the
contrast and power spectrum were statistically com-
parable between animals and man-made objects (ps .
0.29); however, the comparisons between animals and
fruits/vegetables, and between man-made objects and
fruits/vegetables, were significantly different (ps ,
0.01). In Experiment 1B, all images were further
processed with the SHINE toolbox (Willenbockel et al.,
2010; see Figure 1B) to balance low-level visual

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 Journal of Vision (2019) 19(12):22, 1–14
Figure 3. Schematic sample displays with a set size of 6 for the
target present–distractor absent, target present–distractor
present, target absent–distractor absent, and target absent–
distractor present conditions for Experiment 1A. Half of the
participants were asked to search for animals, the other half
were asked to search for man-made objects.
properties including mean luminance, contrast, and
power spectrum (averaged across orientations at each
spatial frequency) across images of all categories.
Procedure
The experiments were run on MATLAB (Math-
Works, Natick, MA) with Psychtoolbox (Brainard,
1997; Kleiner et al., 2007). On each trial, a fixation was
presented at the center of the screen for 500 ms,
followed by a display of 3, 6, or 9 items presented
around an invisible circle (Figure 3). Participants were
asked to determine whether a target was present or
absent by pressing either of two keys on a keyboard as
accurately and as fast as possible. The importance of
accuracy was more emphasized than response speed.
The stimulus display was shown until a response was
made. The distance from the center of each item to the
center of the screen was 5.58, and the visual angle of
each item was 3.68 3 3.68. A chinrest was used to ensure
that the visual distance was fixed to be 57 cm
throughout the study.
Participants were randomly assigned to the animal
or man-made object search group. All participants were
explicitly instructed to search for any items from the
category of interest (e.g., animals for the animal search
group), but there was no mention about the identities
of other two categories (e.g., man-made objects and
fruits/vegetables for the animal search group). In this
way, we were able to examine the effect of attentional
capture from either of animal or man-made object
distractors, which appeared in half of the trials, without
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He & Cheung 5
explicitly reminding participants that three distinct
categories of items would be shown in the study.
For each target-present trial, one item from the target
category was presented (e.g., a squirrel for the animal
search group). For each distractor-present trial, one item
from the distractor category (e.g., a kitchen knife for the
animal search group) was presented. The rest of the items
on the displays were fruits/vegetables. Half of the trials
showed image arrays of round stimuli, and the other half
presented image arrays of elongated stimuli. There were
864 trials in total, with 72 trials in each Target-Present 3
Distractor-Present 3 Set Size condition.
The selection of items on each display and the trial
presentation order were randomized across partici-
pants, and the randomization was matched between
groups in each experiment so that participants search-
ing for either animal or man-made object targets would
see the same displays. The amount of times that the
targets appeared on each of the three, six, or nine
possible locations on the displays were evenly distrib-
uted, while the locations for the appearance of other
items were randomized. Because each animal or man-
made object target and distractor exemplar was
presented only once for each set size during the
experiment, to minimize the possibility that the rarity
of either animals or tools as distractors compared with
fruits/vegetables might account for any distractor
effects, a subset of images of fruits/vegetables (6 out of
14 items from each of the elongated or round sets) were
randomly selected to appear for the same number of
times as the animal/tool distractors. For these items, a
total of 12 out of the 16 exemplars were randomly
selected to maintain identical presentation frequencies
as the animal or man-made object distractors for each
participant, whereas all 16 exemplars were used for all
remaining fruit/vegetable items.
Results
The mean correct response time (RT) for the target
present trials is illustrated in Figure 4, and sensitivity
(d 0 ) for all trials is shown in Table 1. Correct RT was
the main measure here as it reflected the performance
for target present trials, whereas d 0 demonstrated the
performance for both target-present and target-absent
trials. For each experiment, trial outliers with RT
below 150 ms or above three standard deviations of the
average of the individual RT were excluded from the
analyses (average 2.1% of the trials). Two three-way
ANOVAs were conducted on correct RT for target
present trials and d 0 for all trials, with a between-
subjects factor target category (animals vs. man-made
objects), and two within-subject factors, distractor
presence (present vs. absent) and set size (three vs. six
vs. nine).
 Journal of Vision (2019) 19(12):22, 1–14 He & Cheung 6

Figure 4. Mean correct response times on target present trials
averaged across items, as a function of target category,
distractor presence, and set size for Experiments 1A and 1B
(bold lines). Error bars represent the 95% confidence intervals
of the within-subject interaction between distractor presence
and set size. Mean correct response times on target-present
trials for individual items, averaged across distractor-present
and distractor-absent trials, are also shown.
Experiment 1A
For correct RT on target present trials, the main
effect of Target category was significant, F(1, 26) ¼ 9.0,
p ¼ 0.006, g2p ¼ 0.26, with animal search faster than
man-made object search, indicating an overall advan-
tage for animal search. The significant main effect of
distractor presence, F(1, 26) ¼ 6.0, p ¼ 0.021, g2p ¼ 0.19,
revealed longer RT when the distractors were present
relative to when they were absent. The significant main
effect of set size, F(2, 52) ¼ 138.4, p , 0.0001, g2p ¼ 0.84,
showed that RT increased with increased set sizes
(Bonferroni-corrected pairwise comparisons: ps ,
0.0001). None of the interactions was significant
(Target Category 3 Distractor Presence: F(1, 26) ¼ 1.8,
p ¼ 0.19, g2p ¼ 0.07; Target Category 3 Set Size: F(2, 52)
¼ 0.4, p ¼ 0.70, g2p ¼ 0.01; Distractor Presence 3 Set Size:
F(2, 52) ¼ 1.3, p ¼ 0.28, g2p ¼ 0.05; three-way interaction:
F(2, 52) ¼ 0.2, p ¼ 0.82, g2p ¼ 0.01). The search slopes
were comparable between animal search (23.8 ms/item
and 20.6 ms/item with and without the presence of
man-made object distractors), and for man-made
object search (25.4 ms/item and 23.7 ms/item with or
without the presence of animal distractors).
To examine the overall sensitivity performance on
the task, we also calculated d 0 on all trials. Only the
main effect of set size was significant, F(2, 52) ¼ 8.2, p ¼
0.001, g2p ¼ 0.24, with lower d 0 for Set Size 9 compared
with Set Sizes 3 and 6 (Bonferroni-corrected ps , 0.01),
and no statistical difference between set sizes 3 and 6 (p
¼ 0.99). All other main effects or interactions were not
significant (target category: F(1, 26) ¼ 0.01, p ¼ 0.92, g2p
, 0.0001; distractor presence: F(1, 26) ¼ 0.05, p ¼ 0.83,
g2p ¼ 0.002; Target Category 3 Distractor Presence: F(1,
26) ¼ 3.1, p ¼ 0.092, g2p ¼ 0.11; Target Category 3 Set
Size: F(2, 52) ¼ 2.1, p ¼ 0.13, g2p ¼ 0.08; Distractor
Presence 3 Set Size: F(2, 52) ¼ 0.5, p ¼ 0.60, g2p ¼ 0.02;
three-way interaction: F(2, 52) ¼ 0.2, p ¼ 0.79, g2p ¼
0.01).
Experiment 1B
Replicating Experiment 1A, for correct RT on target
present trials, the main effect of target category was

Animal targets Object targets

Set size 3 6 9 3 6 9
Experiment 1A
Distractor present 3.85 (0.49) 3.65 (0.47) 3.47 (0.65) 3.83 (0.56) 3.78 (0.60) 3.66 (0.56)
Distractor absent 3.87 (0.57) 3.88 (0.58) 3.51 (0.51) 3.68 (0.56) 3.73 (0.58) 3.63 (0.48)
Experiment 1B
Distractor present 3.30 (0.79) 3.19 (0.58) 3.19 (0.76) 3.47 (0.61) 3.35 (0.45) 3.28 (0.63)
Distractor absent 3.33 (0.70) 3.14 (0.56) 2.90 (0.80) 3.47 (0.44) 3.40 (0.57) 3.08 (0.56)
Table 1. Mean sensitivity (d 0 ) and standard deviations (in parentheses) as a function of target category, distractor presence, and set
size for Experiments 1A and 1B.

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 Journal of Vision (2019) 19(12):22, 1–14 He & Cheung
significant, F(1, 26) ¼ 8.3, p ¼ 0.008, g2p ¼ 0.24, with
animal search faster than man-made object search. The
main effect of set size was also significant, F(1.43,
37.07) ¼ 91.4, p , 0.0001, g2p ¼ 0.78, showing longer
search time with increased set sizes (Bonferroni-
corrected ps , 0.0001). The Target Category 3 Set Size
interaction approached significance, F(2, 52) ¼ 2.6, p ¼
0.08, g2p ¼ 0.09, but all other main effects or interactions
were not significant (distractor presence: F(1, 26) ¼ 0.5,
p ¼ 0.48, g2p ¼ 0.02; Target Category 3 Distractor
Presence: F(1, 26) ¼ 0.5, p ¼ 0.48, g2p ¼ 0.02; Distractor
Presence 3 Set Size: F(2, 52) ¼ 0.1, p ¼ 0.92, g2p ¼ 0.003;
three-way interaction: F(2, 52) ¼ 2.0, p ¼ 0.15, g2p ¼
0.07). The search slopes for animal targets were 24.0
ms/item with man-made object distractors, and 24.1
ms/item without such distractors. The search slopes for
man-made object targets were 35.0 ms/item with the
presence of animal distractors, and 32.8 ms/item
without animal distractors.
For d 0 , only the main effect of set size was significant,
F(2, 52) ¼ 6.7, p ¼ 0.002, g2p ¼ 0.21, with significantly
lower d 0 for Set Size 9 compared with Set Size 3
(Bonferroni-corrected p , 0.01), and no statistical
difference between other set sizes (ps . 0.10). All other
main effects or interactions were not significant: target
category, F(1, 26) ¼ 0.7, p ¼ 0.41, g2p ¼ 0.03; distractor
presence, F(1, 26) ¼ 2.0, p ¼ 0.17, g2p ¼ 0.07; Target
Category 3 Distractor Presence, F(1, 26) ¼ 0.2, p ¼ 0.65,
g2p ¼ 0.01; Target Category 3 Set Size, F(2, 52) ¼ 0.1, p ¼
0.87, g2p ¼ 0.01; Distractor Presence 3 Set Size, F(2, 52)
¼ 2.8, p ¼ 0.07, g2p ¼ 0.10; three-way interaction: F(2, 52)
¼ 0.2, p ¼ 0.84, g2p ¼ 0.01.
Discussion
With images of comparable visual shape and gist
statistics, we found in both Experiments 1A and 1B
that the search for animal targets remained significantly
faster than that for man-made object targets across set
sizes, suggesting that the search advantage for animals
is not merely driven by visual shape and gist statistics.
Moreover, Experiment 1B further showed that the
search advantage for animals cannot be accounted for
by low-level visual features.
The presence of distractors appeared to attract
attention from searching for the targets in Experiment
1A, suggesting that the animal or tool distractors might
be processed differently from the fruit/vegetable
distractors. However, this effect was not found in
Experiment 1B when the low-level visual properties
were equated among the categories. It is possible that
the target search in Experiment 1A might have been
slowed by the presence of an animal or man-made
object distractor because of the low-level visual
differences between animals or man-made object
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7
distractors, compared with fruit/vegetable distractors.
Alternatively, the effect of actively searching for animal
versus man-made object targets, which was observed in
both Experiments 1A and 1B, appeared to be highly
robust and independent of low-level visual differences,
compared with the stimulus-driven distractor effect.
While it is unlikely that the faster search perfor-
mance for animal than man-made object targets is
solely due to low- and mid-level visual differences
among the categories, one possible account for such
difference in performance is that the categorization
process based on the visual features beyond low- and
mid-level differences is more efficient for animals than
man-made objects. In Experiments 2A and 2B, we
aimed to extend the findings to conditions when
participants were either explicitly told the names of all
target items prior to the search task to minimize the
range of possible targets, or when they were only asked
to detect any items that were not fruits/vegetables and
were not explicitly asked to search for either animal or
man-made object targets. Furthermore, while different
groups of participants searched for either animal or
man-made object targets in Experiments 1A and 1B,
Experiments 2A and 2B further tested whether search
performance would remain faster for animals than
man-made objects by manipulating target category as a
within-subjects factor.
Experiments 2A and 2B
Method
Participants
Forty undergraduate students aged between 18 and
23 of years (mean ¼ 19.8, SD ¼ 1.1, 23 women and 17
men) from the same participant pool as in Experiments
1A and 1B took part in this study; 20 participants
completed Experiment 2A and 20 participants com-
pleted Experiment 2B.
Stimuli and procedure
The stimuli and procedure of Experiments 2A and
2B were identical to those in Experiment 1B, except for
the following changes: The distractor-present trials
were removed. In Experiment 2A, each participant
searched for both animal and man-made object targets
in separate blocks, with the task order counter-
balanced across participants (i.e., half of the partici-
pants searched for animal targets first, and the other
half searched for man-made object targets first).
Participants studied a list of target names (12 animals
or 12 man-made objects) for one minute prior to each
task. In Experiment 2B, participants were asked to
 Journal of Vision (2019) 19(12):22, 1–14
Figure 5. Mean correct response times on target-present trials
averaged across items, as a function of target category and set
size for Experiments 2A and 2B (bold lines). Error bars represent
the 95% confidence intervals of the within-subject interaction
of target category and set size. Mean correct response times on
target-present trials for individual items are also shown.
detect any non-fruit/vegetable item on each trial, and
there was no mention of animals or man-made objects.
The presentation order of animal target trials and man-
made object target trials was randomized.
Animal targets
Set size 3 6
Experiment 2A 3.31 (0.55) 3.30 (0.58)
Experiment 2B 3.54 (0.57) 3.21 (0.52)
Table 2. Mean sensitivity (d 0 ) and standard deviations (in parentheses) as a function of target category and set size for Experiments 2A
and 2B.
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He & Cheung 8
Results
The mean correct response time (RT) for the target
present trials is illustrated in Figure 5, and sensitivity
(d 0 ) for all trials is shown in Table 2. As in Experiment
1A and 1B, trial outliers with RT below 150 ms or
above three standard deviations of the average of the
individual RT for each participant were excluded from
the analyses (average 2.2% of the trials). Two two-way
ANOVAs were conducted on correct RT for target
present trials and d 0 for all trials, with two within-
subject factors target category (animals vs. man-made
objects) and set size (three vs. six vs. nine).
Experiment 2A
For correct RT on target present trials, the main
effect of target category was significant, F(1, 19) ¼ 8.0, p
¼ 0.011, g2p ¼ 0.30, with animal search faster than man-
made object search. The main effect of set size was also
significant, F(2, 38) ¼ 117.5, p , 0.0001, g2p ¼ 0.86,
showing slower search with increased set sizes (Bon-
ferroni-corrected ps , 0.0001). The Target Category 3
Set Size interaction was not significant, F(2, 38) ¼ 1.1, p
¼ 0.35, g2p ¼ 0.05, with 33.1 ms/item for the search slope
for animals and 33.3 ms/item for man-made objects.
For d 0 , the main effect of set size was significant, F(2,
38) ¼ 4.1, p ¼ 0.025, g2p ¼ 0.18, with significantly lower d 0
for Set Size 9 compared with Set Size 3 (Bonferroni-
corrected p , 0.05), and no statistical difference
between other set sizes (ps . 0.10). There was no
significant main effect of target category, F(1, 19) ¼ 2.3,
p ¼ 0.15, g2p ¼ 0.11, or interaction, F(2, 38) ¼ 0.5, p ¼
0.59, g2p ¼ 0.03.
Experiment 2B
For correct RT on target present trials, the main
effect of target category was significant, F(1, 19) ¼ 9.5, p
¼ 0.006, g2p ¼ 0.33, with animal search faster than man-
made object search. The main effect of set size was also
significant, F(2, 38) ¼ 51.5, p , 0.0001, g2p ¼ 0.73,
showing longer search time with increased set sizes
(Bonferroni-corrected ps , 0.0001). The Target Cate-
gory 3 Set Size interaction was not significant, F(2, 38)
¼ 0.5, p ¼ 0.60, g2p ¼ 0.03, revealing no statistical
Object targets
9 3 6 9
3.14 (0.64) 3.20 (0.55) 3.15 (0.68) 2.90 (0.81)
3.11 (0.60) 3.06 (0.47) 2.80 (0.65) 2.66 (0.58)
 Journal of Vision (2019) 19(12):22, 1–14 He & Cheung
difference in the search slopes between animals (34.3
ms/item) and man-made objects (40.1 ms/item).
For d 0 , the main effect of target category was
significant, F(1, 19) ¼ 22.7, p ¼ 0.0001, g2p ¼ 0.54, with
the sensitivity for animal search higher than man-made
object search. The main effect of set size was also
significant, F(2, 38) ¼ 14.8, p , 0.0001, g2p ¼ 0.44, with
significantly higher d 0 for Set Size 3 compared with
both Set Size 6 and Set Size 9 (Bonferroni-corrected ps
, 0.01), and no statistical difference between Set Size 6
and Set Size 9 (p ¼ 0.33). There was no significant
interaction, F(2, 38) ¼ 0.1, p ¼ 0.90, g2p ¼ 0.01).
Discussion
Experiments 2A and 2B replicated the results of both
Experiments 1A and 1B by showing that searching for
animal targets is faster than searching for man-made
objects, even when visual shape, gist statistics, and
other low-level visual features across categories were
comparable. Experiments 2A and 2B further demon-
strated that this category effect was consistently
observed in various conditions, such as when explicit
prior knowledge was given about the specific target
items for the animal or man-made object categories, or
when there was no explicit mention regarding animal or
man-made object categories.
General discussion
Across four experiments that used images of
comparable visual shape and gist statistics across
categories, our findings extended from previous re-
search that used a variety of animal and man-made
object images with naturally varied visual features
within and across categories (e.g., Jackson & Calvillo,
2013; Levin et al., 2001; Long et al., 2017) to show that
the nature of representations that guide visual search
performance may not solely reflect low- or mid-level
visual influences. Specifically, while visual differences
such as curvilinearity and gist statistics clearly have an
important role to support visual categorization (Long
et al., 2017; Loschky & Larson, 2010; Rice et al., 2014;
Zachariou et al., 2018), our findings suggest that these
visual information, in addition to other low-level visual
features such as luminance, contrast, and power
spectrum, cannot entirely account for the faster search
performance for animals than man-made objects.
Although it is likely impossible to rule out all visual
differences between animals and man-made objects,
our stimulus set was designed to rule out a large set of
possible visual features including low-level features,
curvilinearity, elongation, and gist statistics; the com-
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9
parable gist statistics also suggested similar levels of
visual details in the images across the categories. Thus,
the present work highly constrains the scope of
potential explanations as to the visual differences
between categories. Instead, higher-level cognitive
processing, such as the process of categorization, which
involves the interpretation or semantic processing of
the visual features of animals or man-made objects,
may contribute to the category differences and facilitate
visual search performance for animals over man-made
objects. These possibilities are elaborated below.
It is possible that correct categorization of the
animals and man-made objects reflects the different
interpretations of the visual input. Even though the
visual shape and gist statistics are comparable between
categories in this study, there are other available visual
features that observers may utilize to interpret the
meanings of the visual input for categorization.
Previous studies have shown that the interpretation of
ambiguous visual stimuli can shape neural representa-
tions in the occipitotemporal cortex. For instance, with
the absence of actual facial features, face-selective
activations in the fusiform ‘‘face’’ area (FFA) were
observed when the non-face images were interpreted as
faces (Cox, Meyers, & Sinha, 2004; Hadjikhan,
Kveraga, Naik, & Ahlfors, 2009; Summerfield, Egner,
Mangels, & Hirsch, 2006). Nonetheless, successful
categorization of stimuli often relies on correct
interpretations of diagnostic information from the
visual input. What kinds of diagnostic information may
lead us to interpret an image as an animal or a man-
made object? One possibility is that the binding or
configuration of certain features that relies on prior
knowledge about the categories may play a critical role.
For instance, while detecting curvilinear features alone
may likely bias observers to interpret a visual input as
an animal (Long et al., 2017; Zachariou et al., 2018),
various man-made objects such as several of those used
in the current study may also have curvilinear features.
In addition to curvilinear features, detecting an
ensemble of a head, a body, and four limbs may further
facilitate the categorization (Delorme, Richard, &
Fabre-Thorpe, 2010).
It is also important to note that the current study
used a relatively diverse set of animals and man-made
objects (see Appendix); thus, the results were unlikely
due to particular characteristics of a small subset of
animals or objects, but were more likely due to general
knowledge about animals and man-made objects
(Diesendruck & Gelman, 1999; Humphreys et al., 1988;
Laws & Neve, 1999; Patterson, Nestor, & Rogers, 2007;
Taylor, Devereux, Acres, Randall, & Tyler, 2012).
Specifically, the items used for each category in the
experiments were highly different from each other in
terms of diagnostic visual features (e.g., a squirrel vs. a
dolphin). Moreover, the exemplars for each item also
 Journal of Vision (2019) 19(12):22, 1–14 He & Cheung
varied in visual appearance. To successfully categorize
the target items as either animals or man-made objects,
participants would need to utilize information beyond
any visual features that were specific to only a few items
or exemplars of a category. Therefore, the use of our
stimulus set provided further support that our findings
are likely related to general knowledge about possible
visual features of the categories.
Although the current study was not designed to
examine the effects of individual items, the results
showed a range of item differences (Figures 4 and 5).
For animals, a continuum of animacy representation
ranged from mammals to insects have been observed
from lateral to medial occipitotemporal cortex (Con-
nolly et al., 2012; Sha et al., 2015). However, it appears
unlikely that the search performance among different
animals was simply explained by the animacy contin-
uum, as a follow-up observation revealed that the
mammals (e.g., squirrels, mice, dolphins) were not
necessarily among the fastest search, and the insects
(e.g., snails, locusts) were also not necessarily among
the slowest search. Similarly, as man-made objects also
differ in many aspects such as manipulability, size,
portability, and context (Bar, 2004; Mullally &
Maguire, 2011; Peelen & Caramazza, 2012), it remains
unclear which aspects determine the search perfor-
mance for these items. Future work should examine
how different visual and conceptual features of
individual items contribute to the category effects. Our
work provides a method to address these questions.
While the search advantage for animal targets was
robust and was replicated across four experiments, the
extent to which the animal or man-made object
distractors captured attention was less clear, as the
distractor effect was only observed in Experiment 1A,
but it was diminished in Experiment 1B. We speculated
that the animal or man-made object distractors might
have captured attention because of the different image
contrasts or power spectrum compared with neighbor-
ing fruits/vegetables items. The failure to observe
reliable distractor effects in Experiment 1B suggests
that such effects might primarily be influenced by low-
level visual properties, and that different underlying
attentional mechanisms may be involved for the active
search for a target and the involuntary interference
during an active search by a distractor (Langton et al.,
2008; Theeuwes, 1991, 1992; Yantis, 1993).
In contrast with previous studies where visual
features were not strictly controlled (Jackson &
Calvillo, 2013; Levin et al., 2001), we found relatively
steep search slopes for animal and man-made object
categories, and that the search efficiency was compa-
rable for the two categories. These results suggest that
the differential search performance for the two
categories in the current study is unlikely to have
occurred during the pre-attentive stage, as often
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10
indicated by relatively shallow search slopes (Duncan &
Humphreys, 1989; Theeuwes, 1993; Treisman &
Gelade, 1980; Wolfe & Horowitz, 2004). Instead, the
advantage for animal search observed in this study may
arise from a subsequent stage of recognition or
categorization. Nonetheless, it is possible that the
natural variations in visual differences between animals
and man-made objects might have contributed in
determining search efficiency for the categories in the
previous studies (Jackson & Calvillo, 2013; Levin et al.,
2001). Future work should address how the processing
of different sources of information, such as different
levels of visual and semantic features about animals
and man-made objects, may contribute to the differ-
ential performance between the categories in visual
search. Nonetheless, our work adds to a growing
understanding that semantic information may play a
role in visual perception (Coren & Enns, 1993; Lupyan,
2015; Lupyan & Ward, 2013).
To sum up, using images of comparable visual shape
and gist statistics across categories, the current study
provides convergent evidence of faster search for
animals compared with man-made objects. While
category-selective effects between animals and man-
made objects are likely influenced by the vast visual
differences between the categories, as revealed by
previous studies (Long et al., 2017; Zachariou et al.,
2018), we suggest that a focus purely on visual
differences is incomplete. Rather, a full understanding
of category-selective effects in visual perception may
need to incorporate the contributions of high-level
processes, such as categorization processes or semantic
influences, on category-selective effects. More broadly,
the current finding sheds light on the understanding of
the close interactions between perceptual and semantic
systems in human cognition.
Keywords: category selectivity, visual search,
animacy, gist statistics, categorization
Acknowledgments
We thank Emma Wei Chen for helpful discussions,
Daryl Fougnie and Garry Kong for comments on the
manuscript, and Anna Noer for assistance in data
collection. Declarations of interest: none.
Commercial relationships: none.
Corresponding author: Olivia S. Cheung.
Email: olivia.cheung@nyu.edu.
Address: Department of Psychology, Division of
Science, New York University Abu Dhabi, United
Arab Emirates.
 Journal of Vision (2019) 19(12):22, 1–14 He & Cheung
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Appendix
An exemplar of each of the three categories (animals,
man-made objects, fruits/vegetables) is shown in Figure
A1.

Figure A1. An exemplar of each of the three categories: animals, man-made objects, and fruits/vegetables.

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