Content uploaded by Daniela Aguilar
Author content
All content in this area was uploaded by Daniela Aguilar on Feb 03, 2025
Content may be subject to copyright.
AguilarRamirezand R.Gonzalez
Biology of Sex Dierences (2025) 16:6
https://doi.org/10.1186/s13293-025-00693-9
RESEARCH
Sex andhand dierences inhaptic
processing: implications formental rotation
ability
Daniela E. Aguilar Ramirez1* and Claudia L. R. Gonzalez1
Abstract
It has been proposed that the sensorimotor system provides a foundation for the development of cognitive abili-
ties and their hemispheric specialization. In this study, we investigated the potential relationship between haptic
processing and mental rotation ability, both of which are typically lateralized to the right hemisphere. Previous
research has also indicated that males tend to outperform females in both functions. The current study investigates
how the sensorimotor-haptic system relates to mental rotation ability, specifically to examine the influence of hand
performance (as a proxy for hemispheric specialization) and biological sex on this relationship. Seventy-five partici-
pants (n = 41 females) completed a haptic task, and the well-known mental rotation test (MRT) developed by Shepard
and Metzler (Science 171:701–3, 1971). Results confirmed a positive correlation between performance on the haptic
and MRT tasks. Further, males outperformed females in both tasks. However, when sex and hand performance were
considered, males were better in the haptic task, but only when using their left-hand. Moreover, left-hand haptic per-
formance was the sole predictor of MRT performance. These findings suggest that sex differences in haptic process-
ing may contribute to the observed sex differences in mental rotation ability, supporting the view that sensorimotor
processes shape cognitive function and its hemispheric lateralization.
Highlights
• The study of the relationship between the haptic-sensorimotor system and mental rotation ability has remained
scarce.
• Studies have not examined how sex and hand performance contribute to the haptic and mental rotation rela-
tionship.
• Results showed that sex and hand performance were important factors influencing the relationship.
• The sensorimotor system may contribute to the sex differences in mental rotation ability.
• The sensorimotor system appears to help shape cognitive functions and their hemispheric lateralization.
Keywords Haptic processing, Mental rotation, Sex differences, Sensorimotor system, Hemispheric specialization
Open Access
© The Author(s) 2025. Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which
permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the
original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or
other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line
to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory
regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this
licence, visit http:// creat iveco mmons. org/ licen ses/ by/4. 0/. The Creative Commons Public Domain Dedication waiver (http:// creat iveco
mmons. org/ publi cdoma in/ zero/1. 0/) applies to the data made available in this article, unless otherwise stated in a credit line to the data.
Biology of Sex Differences
*Correspondence:
Daniela E. Aguilar Ramirez
d.aguilarramirez@uleth.ca
Page 2 of 9
AguilarRamirezand R.Gonzalez Biology of Sex Dierences (2025) 16:6
Introduction
e embodied cognition theory suggests that sensorimo-
tor experiences shape human perception and interac-
tion with the world [1–5]. Cognitive processes develop
through early sensory and motor interactions, with evi-
dence showing that the same sensorimotor systems used
for physical actions are also involved in mental tasks
like retrieving information [6, 7]. For instance, mental
rotation—a cognitive ability to manipulate objects in the
mind—activates brain areas like the premotor and pri-
mary motor cortices, which are responsible for planning
and executing movements [8, 9]. As Kolb and Whishaw
[10] noted, "mental manipulation is an elaboration of the
neural control of actual manipulation." Exploring the link
between the sensorimotor system and cognition is vital
for understanding brain development and individual dif-
ferences in cognitive abilities.
Mental rotation andthehaptic sensorimotor system
Research has broadly explored the link between mental
rotation and the motor system, suggesting shared mecha-
nisms [11–14]. For example, motor areas activate during
mental rotation [9], and participants perform faster and
more accurately when motor and mental rotation direc-
tions align [12]. Objects harder to physically rotate are
also harder to mentally rotate [15], and embodied objects
(e.g., gloves) are rotated faster than non-embodied ones
(e.g., houses) [16]. Despite this evidence, little is known
about the link between mental rotation and the haptic
system, which integrates touch (cutaneous input) and
proprioception (kinesthetic input) [17]. e haptic sys-
tem is crucial for daily interactions, particularly in low-
visibility situations (e.g., finding keys in a bag). Some
studies suggest individuals with better mental rotation
skills perform better on haptic tasks [18, 19]. Research
in infants shows that manual exploration is key to devel-
oping mental rotation ability [20, 21], highlighting the
role of the haptic system in cognitive development. e
first goal of the current study was tofurther investigate
the relationship between haptic perception and mental
rotation.
Mental rotation andthehaptic system‑sex dierences
Mental rotation ability exhibits significant and consist-
ent sex differences, with males generally outperforming
females, particularly on the Mental Rotation Test (MRT,
[22]). is pattern has been observed across the lifespan
and in over 50 nations [23–28]. In contrast, research on
sex differences in the haptic system is limited and incon-
clusive [29]. Some evidence suggests a male advantage: in
haptic parallelity tasks, blindfolded males made smaller
alignment errors than females [30–32], and males out-
performed females in a shape–texture similarity judg-
ment task [33]. However, other studies found a female
advantage, such as in identifying changes in the positions
of raised line pictures [34]. Additionally, some research
has reported no sex differences, such as in a haptic ver-
sion of the water level test [35, 36]. ese findings high-
light the scarcity and inconsistency in research on sex
differences in haptic perception. e second goal of this
study was to investigate sex differences in haptic percep-
tion and mental rotation ability.
Mental rotation andthehaptic system – hemispheric
specialization
Research has long shown right hemisphere specialization
for mental rotation [37, 38]. Studies using the Shepard
& Metzler [22] task reveal a left visual field advantage in
accuracy and response time in neurologically intact par-
ticipants [39] and similar findings in commissurotomized
Plain language summary
The sensorimotor system has been proposed to contribute to cognitive development. The haptic system, a key com-
ponent of this system, integrates touch and proprioception (awareness of limb positions and movements) to guide
actions. Notably, the right hemisphere of the brain, which controls the left hand, is specialized for haptic process-
ing. This hemisphere is also crucial for mental rotation (MR)—the cognitive ability to mentally visualize how objects
appear when rotated. Interestingly, males have been found to outperform females in both haptic and MR tasks.
This study aimed to explore the relationship between haptic processing and MR ability, particularly examining
whether this relationship is more pronounced in males. Seventy-five participants (41 females) performed both a hap-
tic and a MR task. Findings revealed a correlation between haptic processing and MR ability; specifically, higher perfor-
mance in one predicted higher performance in the other. Importantly, this link was significant only when the haptic
task was performed with the left hand, highlighting the role of the right hemisphere. Additionally, males outper-
formed females in both tasks, with the male advantage in haptic processing observed exclusively with left-hand use.
These results suggest that the male advantage in haptic processing might contribute to the observed male advan-
tage in MR ability. Overall, the findings support the notion that the sensorimotor system, influenced by hemispheric
specialization, plays a role in shaping cognitive functions.
Page 3 of 9
AguilarRamirezand R.Gonzalez Biology of Sex Dierences (2025) 16:6
patients and those with right (but not left) parietal lobe
lesions [40]. Imaging studies pinpoint the right posterior
parietal cortex as a key neural substrate for mental rota-
tion, with consistent activity observed in the right hemi-
sphere in meta-analyses [9, 41].
e right hemisphere also specializes in haptic percep-
tion. Patients with right hemisphere damage show greater
impairment in haptic tasks, such as the Form Board Test,
compared to those with left hemisphere damage [42].
Commissurotomized patients exhibit a left-hand/right-
hemisphere advantage in tasks requiring organization of
scrambled objects by shape or texture [43]. Other studies
support this advantage in texture discrimination, tactual
maze navigation, and shape recognition [44, 45]. Infants
as young as four months old also show a preference for
left-hand exploration [46]. Imaging studies further con-
firm that haptic shape processing is lateralized to the
right posterior parietal lobule [47, 48]. ese findings
suggest shared neural substrates for haptic processing
and mental rotation. e third aim of this study was to
examine hand differences in haptic perception and their
relationship to mental rotation ability.
To summarize, this study aimed to explore the rela-
tionship between haptic perception and mental rotation
ability, with a particular focus on how sex and the hand
used during haptic manipulation may influence this rela-
tionship. We hypothesized, first, a positive relationship
between haptic processing and mental rotation ability;
second, better performance by males in both functions;
and third, a left-hand advantage in haptic processing with
a stronger positive relationship between left-hand haptic
performance and mental rotation ability.
Methods
Participants
Seventy-five (n = 41 females) healthy right-handed par-
ticipants between the ages of 13 to 25years old took part
on this study. Participants self-reported their handed-
ness, sex, and gender (in our sample, all participants were
cisgender). Participants were recruited through word-of-
mouth, media advertisements, and through the Depart-
ment of Psychology at the University of Lethbridge, using
participant management software (Sona Systems). ose
that were university students, received course credits
for their participation. e experiment was approved by
the University of Lethbridge Human Subject Research
Committee.
Tasks
Participants were given the haptic task (Fig.1) and the
MRT (Fig.2). e haptic task consisted of a total of 12
trials, six trials were done with the left-hand and six tri-
als with the right- hand. e order of the trials was ran-
domized within participants. e order of the starting
Fig. 1 The Haptic Task. a The set up- participant picking from a bowl with 12 distinct Lego blocks, the two that they thought made up the model. b
One 2-piece LEGO model
Fig. 2 The Mental Rotation Test. One trial of the Mental Rotation Test (MRT). The target figure on the left was compared to the four figures
on the right. The participant attempted to identify the two figures that were rotated versions of the target. The second and third figures matched
the target figure in this example
Page 4 of 9
AguilarRamirezand R.Gonzalez Biology of Sex Dierences (2025) 16:6
hand (left or right) was also counterbalanced between
participants. Each trial consisted of blindfolded partici-
pant’s haptically exploring a simple 2-piece LEGO model
(Fig.1) for eight seconds with one hand. e Lego model
was made out of two distinct pieces. Each of the 12 mod-
els were different. Immediately after the haptic explora-
tion, the two pieces were placed in a bowl that contained
10 other unique pieces. e participant’s job was to hap-
tically search inside the bowl for the two LEGO pieces
that the made up the model they had just explored. To
be clear, inside the bowl, the two pieces that made up the
model were among twelve pieces; thus, ten pieces were
distractors. e MRT consisted of two sets of 12 trials.
e stimuli for each trial comprised a target figure on
the left and four figures on the right (Fig.2). e partici-
pant’s task was to identify the two figures that matched
the target.
Procedure
Participants were first asked to read and sign a consent
form. en participants were seated at a table (Fig. 1).
For the haptic task participants were told they would be
blindfolded, that they would feel a model with two LEGO
pieces for a few seconds; they were then instructed to
search for the two pieces that made up the model. ey
were told that they were only allowed to use one hand
(either left or right) to feel the model and find the pieces,
and that they could not use both hands. Furthermore,
they were told that once they found the pieces, they
would take them out of the bowl and locate them on the
side, on top of the table. ey were asked to do the task
as accurate and as fast as they could. e experimenter
then handed the participant a blindfold to cover their
eyes. e bowl containing the twelve pieces was placed
on top of the table, in front of the participant. e experi-
menter then placed the model (two pieces together) on
the participants’ corresponding hand. e participants
felt the model for eight seconds, starting when the exper-
imenter placed the model on the participants’ hand and
ending when the experimenter said the time was up. e
experimenter then placed the two pieces inside the bowl
containing the 10 distractors and started the stopwatch
as soon as the participant began searching for the pieces
inside the bowl. e experimenter stopped the stopwatch
once the participant had placed the two pieces outside of
the bowl and on the table. Each of the trials followed the
same procedure. Following the haptic task, the MRT was
given to the participants (Fig.2). For the MRT, partici-
pants were instructed to choose the two stimuli out of the
four options that matched the target stimuli. Participants
were given a 3-min limit to complete each of the two sets
with a 3-min break between sets (Peters etal. 1995).
Data analysis
For the haptic task, there were two dependent vari-
ables: errors and time to complete the task. An error
was recorded if the participant took out of the bowl
a piece that did not match any of the two pieces that
made up the model. e time taken to find the pieces
inside each of the bowls (i.e. each trial) was recorded.
e MRT was scored by taking the sum of only the tri-
als in which the two answers were correct, dividing by
the maximum score of 24, and then multiplying by 100.
Pearson correlation coefficients were calculated
between the haptic task time, haptic errors, and MRT
to examine the relationship between the haptic task and
MRT performance. A mixed design ANOVA was used
to investigate sex differences in the haptic task, and the
MRT with Sex as fixed factor. Furthermore, to explore
Sex and Hand differences in haptic task performance,
a repeated measures ANOVA was used, with Sex as the
between-participant factor and hand as within. Lastly,
to investigate hand differences in haptic perception
and its relationship to mental rotation ability, a linear
regression analysis was conducted to determine if any
of the dependent variables (left-hand errors, right-hand
errors, left-hand time, and right-hand time), was the
predictor of performance in the MRT. IBM SPSSStatis-
tics (Version 29) was used for all analyses. e alpha
level for all comparisons was 0.05.
Results
Haptic processing andMRT relationship
Figure3 shows the results of the correlational analysis.
ere was a significant negative correlation (r = −0.40)
between the number of errors on the haptic task and
MRT performance, the more errors the participants
made, the worse their performance on the MRT. No
significant difference was found for the amount of time
participants took in solving the haptic task.
Sex dierences—haptic task andMRT
e mixed design ANOVA (see Table 1) revealed a
significant main effect of Sex in the number of errors
F (1,73) = 12.19, p < 0.001, η2p = 0.14 and in the time F
(1,73) = 7.62, p < 0.01, η2p = 0.09 participants took in the
haptic task. Males made fewer errors than females and
took less time than females but this was only true for
the left-hand. ere was not a significant effect of Sex
in the errors or time participants took in the haptic task
with their right hand. For the MRT task, a significant
main effect of Sex, F (1,73) = 9.87, p < 0.01, η2p = 0.12 was
found, males had better scores than females (Table1).
Page 5 of 9
AguilarRamirezand R.Gonzalez Biology of Sex Dierences (2025) 16:6
Haptic task‑ sex andhand dierences
Performance on the haptic task was analyzed with Sex
(female, male) by Hand (left, right) repeated measures
ANOVA, with Sex as between-participant factor and
Hand as a within-participant factor. Regarding haptic
task errors, a significant main effect of Sex was found
F (1,73) = 8.49, p < 0.01, η2p = 0.07 and a significant Sex
by Hand interaction F (1,73) = 4.09, p < 0.05, η2p = 0.02.
Males made fewer errors than females when using their
left-hand but not when using their right hand (Fig.4A).
Regarding haptic task times, a significant main effect
of Sex was found F (1,73) = 4.09, p < 0.05, η2p = 0.04; w ith
Fig. 3 Results of the correlation analysis for haptic task and MRT performance
Table 1 Means and standard errors for the dependent variables. Please note the sex difference when using the left, but not the right
hand
Dependent variable Grand mean Female participants Male participants F statistic
Haptic Left Errors (#) 1.76 ± 0.20 2.34 ± 0.29 1.06 ± 0.21 F (1,73) = 12.19, p < 0.01, η2p = 0.14
Haptic Left Time (s) 31.99 ± 1.49 35.58 ± 2.18 27.65 ± 1.75 F (1,73) = 7.62, p < 0.01, η2p = 0.09
Haptic Right Errors (#) 1.8 ± 0.18 2.00 ± 0.26 1.60 ± 0.26 F (1,73) = 1.34, p > 0.1, η2p = 0.18
Haptic Right Time (s) 32.97 ± 1.56 34.18 ± 2.13 31.51 ± 2.31 F (1,73) = 0.72, p > 0.1, η2p = 0.10
MRT score (%) 43.94 ± 2.61 38.89 ± 3.20 52.45 ± 3.83 F (1,73) = 9.87, p < 0.01, η2p = 0.12
Fig. 4 Haptic Task errors and times for each Hand (left, right) and Sex (female, male). p ≤ 0.01**, p ≤ 0.001***. Error bars represent standard errors
Page 6 of 9
AguilarRamirezand R.Gonzalez Biology of Sex Dierences (2025) 16:6
males completing the task faster. A Sex by Hand inter-
action approached significance F (1,73) = 3.16, p = 0.08,
η2p = 0.01. To further explore this marginal interaction,
t-tests were conducted between male and female partici-
pants’ performance with the right and left-hands. Males
were significantly faster than females when using their
left-hand but not when using their right hand (Fig.4B).
Haptic task andMRT relationship—hand dierences
To further explore the relationship between left- and
right-haptic task performance and mental rotation abil-
ity, a regression analysis was used. e model included
MRT as the dependent variable and left-hand haptic task
errors, right-hand haptic task errors, left-hand haptic
time, and right-hand haptic time as potential predictors
(see Table2). e multiple linear regression analysis was
significant (F (1,73) = 4.49, p < 0.01, R2 = 0.20). e left-
hand number of errors was the sole significant predictor
of MRT performance (p < 0.01).
Discussion
e results of the current investigation revealed a signifi-
cant relationship between errors in the haptic task and
performance on the Mental Rotation Test (MRT): more
errors in the haptic task were associated with poorer
MRT performance, supporting our first hypothesis.
is finding aligns with theembodied cognition theory,
which posits that active physical manipulation of objects
is linked to mental manipulation. Shepard [49] proposed
that mental representations are “isomorphic” to physical
rotation, and previous studies have shown that physical
rotation can hinder or enhance mental rotation depend-
ing on directional alignment [50, 51]. However, these
studies primarily involved tactile stimulation or handheld
sensors, whereas the current study’s haptic task included
active object manipulation, engaging both touch and pro-
prioception, making it more ecologically valid.
Interestingly, no significant relationship was found
between haptic task completion time and MRT per-
formance, although the correlation was in the expected
direction (longer times were associated with worse
MRT scores). is lack of significance may be due to
high variability in task completion times; some partici-
pants may have quickly stumbled upon the target pieces,
while others took longer, independent of their accuracy.
Indeed, the number of errors was not correlated with
task time (r = 0.02), suggesting accuracy and speed are
not directly related in the haptic task. Previous research
has also shown that speed and accuracy do not always
align [52]. While speed measures can provide insights
into cognitive processes, their relationship with accu-
racy is complex and sometimes inconsistent [53]. is
discrepancy might stem from the dual-process theory,
which distinguishes between fast, automatic thinking and
slower, deliberate reasoning [54]. ese two modes of
thinking could explain the variability in how participants
approached the haptic task and theMRT.
Regarding sex differences, results showed that males
made fewer errors in the haptic task and scored better
in the MRT. us, our second hypothesis was supported,
a male advantage was found in both tasks. ese results
are in line with other researchers who have found a male
advantage in haptic perception. ese studies, however,
have mostly used the haptic parallelity task [30–32, 55,
56] where participants aligned a reference bar (previously
placed in a different orientation) to the test bar. e par-
allelity task taps mostly on the ability to orient objects
in space and less on haptic discrimination of shape and/
or texture (as the task used in the current study). To our
knowledge few studies have explored sex differences
in haptic discrimination of shape and/or texture [33,
57], withCohen and Levy (1986) [33], reporting a male
advantage. us, our findings make a significant contri-
bution to the literature by showing a male advantage spe-
cifically for haptic processing of object shape. As well, our
findings add to the large body of literature supporting a
male advantage in mental rotation ability. To our knowl-
edge, this is the first study to show a male advantage in
these two processes in the same sample of participants.
When examining sex and hand performance in the
haptic task, results showed that males outperformed
females by making fewer errors and completing the task
faster, but only when using the left hand. is aligns
with Witelson’s [57] findings, where boys (ages 6–13)
Table 2 Results of the regression analysis
The only signicant predictor of MRT performance was the left-hand errors
Independent variables Unstandardized BCoecients of standard
error Standardized
coecients beta t Sig.
(Constant) 65.69 8.01 8.16 < 0.01
Left‑hand errors (#) −4.43 1.55 −0.33 −2.86 0.01
Right-hand errors (#) − 2.22 1.69 − 0.15 − 1.32 0.19
Left-hand time (s) − 0.22 0.23 − 0.13 − 0.99 0.33
Right-hand time (s) − 0.08 0.21 − 0.05 − 0.40 0.69
Page 7 of 9
AguilarRamirezand R.Gonzalez Biology of Sex Dierences (2025) 16:6
exhibited a left-hand advantage in a haptic shape dis-
crimination task, though no overall sex differences were
found. Witelson attributed this left-hand advantage to
the right hemisphere’s specialization for spatial pro-
cessing which, Witelson argued, is more pronounced in
males, while females exhibit more bilateral processing.
Subsequent studies (e.g., Nilsson & Geffen [58]) repli-
cated the male left-hand advantage and similarly linked
it to right hemisphere spatial strategies, contrasting with
females’ likely use of bilateral cognitive strategies. is
male bias toward right hemisphere dominance in spatial
tasks, including mental rotation [59, 60], may explain the
left-hand advantage observed in haptic tasks. Support-
ing this view, clinical and imaging studies suggest males
show stronger hemispheric specialization, while females
exhibit greater inter-hemispheric connectivity [61, 62].
However, other research has failed to replicate these
findings (e.g., Cranney & Ashton [63]) or has reported a
right-hand advantage with no sex differences [64]. ese
inconsistencies highlight the need for further research
exploring the interplay between sex, hand dominance,
and haptic processing to determine whether the left-hand
male advantage stems from spatial processing, haptic
strategies, or an interaction of these factors.
To more directly examine the relationship between
hand performance on the haptic task and the MRT, a
regression analysis was done. Results showed that the
number of errors with the left-hand was the only predic-
tor of MRT performance. us, confirming our hypoth-
esis of a left-hand positive relationship with mental
rotation. is finding expands previous literature of
right hemisphere specialization for haptic processing
and mental rotation ability [9, 48] to demonstrate that
these processes are linked. erefore, the results suggest
that haptic processing, and mental rotation may share
neural substrates which have been identified in the pari-
etal cortex [65, 66]. However, behavioural experiments
that include imaging techniques aimed at exploring this
relationship are missing. Future research should first,
explore if these processes share neural substates; second,
if there is sexual dimorphism in these substrates; and
third, how these brain processes (i.e., haptic and mental
rotation) interact during development. Overall, research
addressing these aspects could provide insights into the
importance of the haptic system for the development of
spatial cognition and the associated sex differences in
these processes.
One limitation of this study lies in the specific nature of
the task. Unlike previous research, which often employed
nonsensical shapes or objects with minimal tactile cues
[48, 54, 58] this study utilized Lego bricks, which may
provide richer haptic information. Future research could
explore a wider variety of objects with varying levels of
haptic complexity to determine whether task difficulty
modulates sex differences in haptic processing.
In conclusion, the results of this study support a close
relationship between haptic processing and mental rota-
tion ability that is influenced by sex and hand. A male
advantage was found in haptic processing and mental
rotation ability. However, the male advantage in haptic
processing appears to exist only for the left hand. Fur-
thermore, only a left-hand relationship was found with
MRT performance. In sum, this study provides evidence
of how the sensorimotor system and cognition are inter-
related, noting that this interrelatedness differs by sex.
Acknowledgements
The authors thank all participants for their contribution to this project. The
authors would like to thank the Natural Sciences and Engineering Research
Council of Canada for supporting this research.
Author contributions
D. A. collected the data, analyzed the data, and wrote the main manuscript
text. C.G developed protocol, guided in data analyses, and helped with manu-
script writing. All authors reviewed the manuscript.
Funding
This project was funded by the Natural Sciences and Engineering Research
Council of Canada with a Tier II Canada Research Chair and a Discovery Grant
awarded to Dr. Claudia Gonzalez (Grant no. 14367).
Data availability
The materials and data that support the findings of this study are available
from the corresponding author upon reasonable request.
Declarations
Ethics approval and consent to participate
All procedures performed in studies involving human participants were in
accordance with the ethical standards of the institutional and/or national
research committee and with the 1964 Helsinki declaration and its later
amendments or comparable ethical standards. Informed consent: Informed
consent was obtained from all individual participants included in the study.
Consent for publication
Informed consent was obtained from all individualparticipants included in the
study.
Competing interests
The authors declare no competing interests.
Author details
1 Department of Kinesiology and Physical Education, University of Lethbridge,
4401 University Drive, Lethbridge, AB T1K 3M4, Canada.
Received: 12 September 2024 Accepted: 25 January 2025
References
1. Clark A, Chalmers D. The extended mind. Analysis. 1998;58:7–19. https://
doi. org/ 10. 1093/ analys/ 58.1.7.
2. Butz MV. Toward a unified sub-symbolic computational theory of cogni-
tion. Front Psychol. 2016. https:// doi. org/ 10. 3389/ fpsyg. 2016. 00925.
3. Tomlinson SP, Davis NJ, Morgan HM, Bracewell RM. Hemispheric speciali-
sation in haptic processing. Neuropsychologia. 2011;49:2703–10. https://
doi. org/ 10. 1016/j. neuro psych ologia. 2011. 05. 018.
Page 8 of 9
AguilarRamirezand R.Gonzalez Biology of Sex Dierences (2025) 16:6
4. Mcglone J. Sex differences in human brain asymmetry: a critical survey.
Behav Brain Sci. 1980;3:215–27.
5. Scheuringer A, Harris T-A, Pletzer B. Recruiting the right hemisphere:
sex differences in inter-hemispheric communication during semantic
verbal fluency. Brain Language. 2020;207:104814. https:// doi. org/ 10.
1016/j. bandl. 2020. 104814.
6. Jeannerod M, Decety J. Mental motor imagery: a window into the
representational stages of action. Curr Opin Neurobiol. 1995;5:727–32.
https:// doi. org/ 10. 1016/ 0959- 4388(95) 80099-9.
7. Oldrati V, Finisguerra A, Avenanti A, Aglioti SM, Urgesi C. Differential
influence of the dorsal premotor and primary somatosensory cortex
on corticospinal excitability during kinesthetic and visual motor
imagery: A low-frequency repetitive transcranial magnetic stimulation
study. Brain Sci. 2021. https:// doi. org/ 10. 3390/ brain sci11 091196.
8. Ganis G, Keenan JP, Kosslyn SM, Pascual-Leone A. Transcranial magnetic
stimulation of primary motor cortex affects mental rotation. Cereb
Cortex. 2000;10:175–80. https:// doi. org/ 10. 1093/ cercor/ 10.2. 175.
9. Zacks JM. Neuroimaging studies of mental rotation: a meta-analysis
and review. J Cogn Neurosci. 2008;20:1–19. https:// doi. org/ 10. 1162/
jocn. 2008. 20013.
10. Kolb IQ, et al. Fundamentals of human neuropsychology. 5th ed. New
York: Worth Publishers; 2003.
11. Voyer D, Jansen P. Motor expertise and performance in spatial tasks: a
meta-analysis. Hum Mov Sci. 2017;54:110–24. https:// doi. org/ 10. 1016/j.
humov. 2017. 04. 004.
12. Wexler M, Kosslyn SM, Berthoz A. Motor processes in mental rotation.
Cognition. 1998;68:77–94. https:// doi. org/ 10. 1016/ S0010- 0277(98)
00032-8.
13. Moreau D, Mansy-Dannay A, Clerc J, Guerrien A. Spatial ability and
motor performance: Assessing mental rotation processes in elite and
novice athletes. Int J Sport Psychol. 2011;42:525–54.
14. Schwarzer G, Freitag C, Schum N. How crawling and manual object
exploration are related to the mental rotation abilities of 9-month-old
infants. Front Psychol. 2013. https:// doi. org/ 10. 3389/ fpsyg. 2013. 00097.
15. Flusberg SJ, Boroditsky L. Are things that are hard to physically move
also hard to imagine moving? Psychon Bull Rev. 2011;18:158–64.
https:// doi. org/ 10. 3758/ s13423- 010- 0024-2.
16. Suggate HS, Lehmann J, Jansen P. Cognition embodied: mental rota-
tion is faster for objects that imply a greater body–object interaction. J
Cogn Psychol. 2019;31:876–90. https:// doi. org/ 10. 1080/ 20445 911. 2019.
16786 27.
17. Lederman SJ, Klatzky RL. Haptic perception: a tutorial. Atten Percept
Psychophys. 2009;71:1439–59. https:// doi. org/ 10. 3758/ APP. 71.7. 1439.
18. Lebaz S, Jouffrais C, Picard D. Haptic identification of raised-line
drawings: high visuospatial imagers outperform low visuospatial
imagers. Psychol Res. 2012;76:667–75. https:// doi. org/ 10. 1007/
s00426- 011- 0351-6.
19. Kalisch J-CAK Tobias, Kattenstroth. Cognitive and tactile factors affect-
ing human haptic performance in later life. PLOS ONE 2012;7:1–11.
https:// doi. org/ 10. 1371/ journ al. pone. 00304 20.
20. Frick A, Möhring W. Mental object rotation and motor development
in 8- and 10-month-old infants. J Exp Child Psychol. 2013;115:708–20.
https:// doi. org/ 10. 1016/j. jecp. 2013. 04. 001.
21. Frick A, Wang S. Mental spatial transformations in 14- and 16-month-
old infants: Effects of action and observational experience. Child Dev.
2014;85:278–93. https:// doi. org/ 10. 1111/ cdev. 12116.
22. Shepard RN, Metzler J. Mental rotation of three-dimensional objects.
Science. 1971;171:701–3. https:// doi. org/ 10. 1126/ scien ce. 171. 3972. 701.
23. Linn MC, Petersen AC. Emergence and characterization of sex differ-
ences in spatial ability: a meta-analysis. Child Dev. 1985;56:1479–98.
24. Voyer D, Voyer S, Bryden MP. Magnitude of sex differences in spatial
abilities: a meta-analysis and consideration of critical variables. Psychol
Bull. 1995;117:250–70. https:// doi. org/ 10. 1037/ 0033- 2909. 117.2. 250.
25. Lippa RA, Collaer ML, Peters M. Sex differences in mental rotation and
line angle judgments are positively associated with gender equal-
ity and economic development across 53 nations. Arch Sex Behav.
2010;39:990–7. https:// doi. org/ 10. 1007/ s10508- 008- 9460-8.
26. Levine SC, Foley A, Lourenco S, Ehrlich S, Ratliff K. Sex differences in
spatial cognition: advancing the conversation. Wiley Interdiscip Rev
Cogn Sci. 2016;7:127–55. https:// doi. org/ 10. 1002/ wcs. 1380.
27. Aguilar Ramirez DE, Blinch J, Gonzalez CLR. One brick at a time: Building a
developmental profile of spatial abilities. Dev Psychobiol. 2021;63:e22155.
https:// doi. org/ 10. 1002/ dev. 22155.
28. Aguilar Ramirez DE, Blinch J, Takeda K, Copeland JL, Gonzalez CLR. Dif-
ferential effects of aging on spatial abilities. Exp Brain Res. 2022;240:1579–
88. https:// doi. org/ 10. 1007/ s00221- 022- 06363-1.
29. Fernandes AM, Albuquerque PB. Tactual perception: a review of experi-
mental variables and procedures. Cogn Process. 2012;13:285–301. https://
doi. org/ 10. 1007/ s10339- 012- 0443-2.
30. Kappers AML. Large systematic deviations in a bimanual parallelity task:
Further analysis of contributing factors. Acta Psychol. 2003;114:131–45.
https:// doi. org/ 10. 1016/ S0001- 6918(03) 00063-5.
31. Hermens F, Kappers AML, Gielen SCAM. The structure of frontoparal-
lel haptic space is task dependent. Percep Psychophys. 2006;68:62–75.
https:// doi. org/ 10. 3758/ BF031 93656.
32. Zuidhoek S, Kappers AML, Postma A. Haptic orientation perception:
Sex differences and lateralization of functions. Neuropsychologia.
2007;45:332–41. https:// doi. org/ 10. 1016/j. neuro psych ologia. 2006. 05. 032.
33. Cohen H, Levy JJ. Sex differences in categorization of tactile stimuli.
Percept Mot Skills. 1986;63:83–6. https:// doi. org/ 10. 2466/ pms. 1986. 63.1.
83.
34. Heller MA, Jones ML, Walk AM, Schnarr R, Hasara A, Litwiller B. Sex differ-
ences in the haptic change task. J Gen Psychol. 2009;137:49–62. https://
doi. org/ 10. 1080/ 00221 30090 32930 63.
35. Berthiaume F, Robert M, St-Onge R, Pelletier J. Absence of a gender
difference in a haptic version of the water-level task. Bull Psychon Soc.
1993;31:57–60. https:// doi. org/ 10. 3758/ BF033 34140.
36. Robert M, Pelletier J, St-Onge R, Berthiaume F. Women’s deficiency in
water-level representation: present in visual conditions yet absent in
haptic contexts. Acta Psychol. 1994;87:19–32. https:// doi. org/ 10. 1016/
0001- 6918(94) 90064-7.
37. Ratcliff G. Spatial thought, mental rotation and the right cerebral hemi-
sphere. Neuropsychologia. 1979;17:49–54. https:// doi. org/ 10. 1016/ 0028-
3932(79) 90021-6.
38. Farah MJ. The neural basis of mental imagery. Trends Neurosci.
1989;12:395–9. https:// doi. org/ 10. 1016/ 0166- 2236(89) 90079-9.
39. Ditunno PL, Mann VA. Right hemisphere specialization for mental rotation
in normals and brain damaged subjects. Cortex. 1990;26:177–88. https://
doi. org/ 10. 1016/ s0010- 9452(13) 80349-8.
40. Corballis MC, Sergent J. Hemispheric specialization for mental rotation.
Cortex. 1989;25:15–25. https:// doi. org/ 10. 1016/ s0010- 9452(89) 80002-4.
41. Harris IM, Egan GF, Sonkkila C, Tochon-Danguy HJ, Paxinos G, Watson JDG.
Selective right parietal lobe activation during mental rotation: a paramet-
ric PET study. Brain. 2000;123:65–73. https:// doi. org/ 10. 1093/ brain/ 123.1.
65.
42. De Renzi E. Nonverbal memory and hemispheric side of lesion. Neuropsy-
chologia. 1968;6:181–9. https:// doi. org/ 10. 1016/ 0028- 3932(68) 90018-3.
43. Sperry RW, Gazzaniga MS, Bogen JE. Interhemispheric relationships: the
neocortical commissures; syndromes of hemisphere disconnection, vol.
4. New York: John Wiley Sons; 1969.
44. Stone KD, Gonzalez CL. The contributions of vision and haptics to reach-
ing and grasping. Front Psychol. 2015. https:// doi. org/ 10. 3389/ fpsyg.
2015. 01403.
45. Fagot J, Lacreuse A, Vauclair J. Haptic discrimination of nonsense shapes:
hand exploratory strategies but not accuracy reveal laterality effects.
Brain Cogn. 1993;21:212–25. https:// doi. org/ 10. 1006/ brcg. 1993. 1017.
46. Morange-Majoux F. Manual exploration of consistency (soft vs hard) and
handedness in infants from 4 to 6 months old. Laterality. 2011;16:292–
312. https:// doi. org/ 10. 1080/ 13576 50090 35536 89.
47. Butler AJ, Fink GR, Dohle C, Wunderlich G, Tellmann L, Seitz RJ, et al.
Neural mechanisms underlying reaching for remembered targets cued
kinesthetically or visually in left or right hemispace. Hum Brain Map.
2004;21:165–77. https:// doi. org/ 10. 1002/ hbm. 20001.
48. Marangon M, Kubiak A, Króliczak G. Haptically guided grasping. fMRI
shows right-hemisphere parietal stimulus encoding, and bilateral dorso-
ventral parietal gradients of object- and action-related processing during
grasp execution. Front Hum Neurosci. 2016. https:// doi. org/ 10. 3389/
fnhum. 2015. 00691.
49. Shepard RN. Form, formation, and transformation of internal representa-
tions. Information processing and cognition. 1st ed. Routledge; 1975.
Page 9 of 9
AguilarRamirezand R.Gonzalez Biology of Sex Dierences (2025) 16:6
50. Gardony AL, Taylor HA, Brunyé TT. What does physical rotation reveal
about mental rotation? Psychol Sci. 2014;25:605–12. https:// doi. org/ 10.
1177/ 09567 97613 503174.
51. Lohmann J, Rolke B, Butz MV. In touch with mental rotation: Interactions
between mental and tactile rotations and motor responses. Exp Brain
Res. 2017;235:1063–79. https:// doi. org/ 10. 1007/ s00221- 016- 4861-8.
52. Lohman DF. Spatial ability: Individual differences in speed and level.
Standford University Press; 1979.
53. Kyllonen PC, Zu J. Use of response time for measuring cognitive ability. J
Intell. 2016. https:// doi. org/ 10. 3390/ jinte llige nce40 40014.
54. Evans JSBT, Stanovich KE. Dual-process theories of higher cognition:
advancing the debate. Perspect Psychol Sci. 2013;8:223–41. https:// doi.
org/ 10. 1177/ 17456 91612 460685.
55. Kappers AM. Haptic space processing–allocentric and egocentric refer-
ence frames. Can J Exp Psychol. 2007;61:208–18. https:// doi. org/ 10. 1037/
cjep2 007022.
56. Van Mier HI. Effects of visual information regarding allocentric processing
in haptic parallelity matching. Acta Psychol. 2013;144:352–60. https:// doi.
org/ 10. 1016/j. actpsy. 2013. 07. 003.
57. Witelson SF. Sex and the single hemisphere: specialization of the right
hemisphere for spatial processing. Science. 1976;193:425–7. https:// doi.
org/ 10. 1126/ scien ce. 935879.
58. Nilsson J, Geffen G. Perception of similarity and laterality effects in tactile
shape recognition. Cortex. 1987;23:599–614. https:// doi. org/ 10. 1016/
S0010- 9452(87) 80051-5.
59. Roberts JE, Ann BM. Two- and three-dimensional mental rotation tasks
lead to different parietal laterality for men and women. Int J Psychophys-
iol. 2003;50:235–46. https:// doi. org/ 10. 1016/ S0167- 8760(03) 00195-8.
60. Vogel JJ, Bowers CA, Vogel DS. Cerebral lateralization of spatial abilities:
a meta-analysis. Brain Cogn. 2003;52:197–204. https:// doi. org/ 10. 1016/
s0278- 2626(03) 00056-3.
61. Rilea SL, Roskos-Ewoldsen B, Boles D. Sex differences in spatial ability: A
lateralization of function approach. Brain Cogn. 2004;56:332–43. https://
doi. org/ 10. 1016/j. bandc. 2004. 09. 002.
62. Ingalhalikar M, Smith A, Parker D, Satterthwaite TD, Elliott MA, Ruparel K,
et al. Sex differences in the structural connectome of the human brain.
Proc Natl Acad Sci U S A. 2014;111:823–8. https:// doi. org/ 10. 1073/ pnas.
13169 09110.
63. Cranney J, Ashton R. Witelson’s dichhaptic task as a measure of hemi-
spheric asymmetry in deaf and hearing populations. Neuropsychologia.
1980;18:95–8. https:// doi. org/ 10. 1016/ 0028- 3932(80) 90089-5.
64. Hannay HJ, Smith AC. Dichhaptic perception of forms by normal adults.
Percept Mot Skills. 1979;49:991–1000. https:// doi. org/ 10. 2466/ pms. 1979.
49.3. 991.
65. Sack AT. Parietal cortex and spatial cognition. Behav Brain Res.
2009;202:153–61. https:// doi. org/ 10. 1016/j. bbr. 2009. 03. 012.
66. Lee Masson H, Bulthé J, Op Beeck HP, et al. Visual and haptic shape
processing in the human brain: unisensory processing, multisensory
convergence, and top-down influences. Cereb Cortex. 2016;26:3402–12.
https:// doi. org/ 10. 1093/ cercor/ bhv170.
Publisher’s Note
Springer Nature remains neutral with regard to jurisdictional claims in pub-
lished maps and institutional affiliations.