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Investigating the Motor Cortex’s (M1) Causational Role in Mental Rotation using

Transcranial Magnetic Stimulation (TMS)

Chloe Allen-Sciberras

Melbourne School of Psychological Sciences, University of Melbourne

Biological Psychology (PSYC20006)

Catherin Maguire

19/04/2024

1,643
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Abstract

The primary motor cortex (M1) is known for directing the body’s skeletal muscles to perform

voluntary movements, however, previous studies have suggested that it may play a role in

cognitive tasks such as the mental rotation of objects. This study used single-pulse transcranial

magnetic stimulation (TMS) to investigate the primary motor cortex’s causal involvement in the

mental rotation of Shepard Figures and hands. 60 participants, aged 18-30, were involved. 30 of

the participants were allocated to the Shepards Figures condition (M = 23.60 years,15 male, 15

female), and 30 (M = 24.03 years, 14 male, 16 female) were assigned to the hands condition. A

paired samples t-test measured the mean difference between reaction times (RT) for the mental

rotation of Shepard Figures and Hands under real and sham TMS conditions. There was an

increase in RT under real TMS for both. These results were considered statistically significant,

with p-values of p<0.001 and p=0.001, respectively. The Shepard Figures condition had a

Cohen’s value of d=0.70 and the hands condition d=0.60, both of which suggest a practical

significance for this study. These findings imply a causal relationship between the M1 and

mental rotation and prompt further investigation into the cognitive applications of the M1.

Keywords: Primary Motor Cortex (M1), mental rotation, Transcranial Magnetic

Stimulation (TMS), Shepard Figures, reaction time (RT)

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Investigating the Motor Cortex’s (M1) Causational Role in Mental Rotation using

Transcranial Magnetic Stimulation (TMS)

Mental rotation is a cognitive process that involves 2D or 3D objects being imaginatively

formed and rotated (Rholetter, 2022). Traditionally, mental rotation is associated with brain

regions responsible for strictly cognitive processes but studies have suggested a correlation

with the primary motor cortex (M1). The M1 is a region at the rear of the frontal lobe that

mediates the planning, initiation and control of voluntary movements (Purves et al., 2001). This

understanding of the M1 made it difficult for researchers to comprehend why it may be involved

in processes that do not require motor neurons (Eisenegger et al., 2007). To answer these

questions, researchers conducted studies using transcranial magnetic stimulation (TMS). TMS

utilises a rapidly changing magnetic field to stimulate the brain and cause neurons to fire

randomly. This alters the ongoing activity of the neural tissue under the TMS coil, creating

‘virtual lesions’ (Harris et al., 2008). Single-pulse TMS is ideal for investigating causal

relationships as it simply investigates brain activity, whereas repetitive TMS may alter brain

activity (Klomjai et al., 2015). The stimulating properties of TMS act as ‘neural noise’, masking

intended brain activity and increase RTs for tasks that require the brain region underneath the

coil. This masking and resultant slower RTs demonstrate a causal relationship.

Kosslyn et al. (1998) used positron emission tomography (PET) to monitor regional

cerebral blood flow (rCBF) whilst participants underwent two mental rotation tasks. One task

involved mentally rotating ‘angular branching forms’ or Shepard Figures (Shepard & Metzler,

1971), and the other involved the mental rotation of hands. A baseline condition was present,

where participants were presented with identical stimuli but were not required to mentally rotate

it. Kosslyn and colleagues discovered that M1 activation was present during the rotation of

hands but not Shepard Figures, this was attributed to the numerous mental rotation techniques.

Suggesting that the M1 would be activated when visualisation of physical manipulation was

used (e.g. imagining turning the object with your hand). When the Shepard Figures were being
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rotated participants would imagine an external force turning them, and would not activate their

M1. Overall, Kosslyn et al. (1998) determined that mental rotation involved two mechanisms,

one that had similar preparation to skeletal movement (internal) and one that did not (external).

Eisenegger and colleagues (2007) used single-pulse TMS on the left M1 to identify

neural activity when participants visualised rotating Shepard Figures (Shepard & Metzler, 1971).

This study used a probing excitability approach by recording motor-evoked potentials (MEPs)

that recognise the activation of motor pathways (Doyal et al., 2022) during mental rotation.

Mental rotation was compared to reading and it produced more brain activity, ruling out previous

ideas that verbal strategies were a solution for mental rotation tasks. Even though asking

participants to imagine rotating objects with their hands aligns with the strategy-dependent

findings of Kosslyn et al. (1998), Eisenegger et al. (2007) insisted that the ‘spill-over effect’ was

the most likely explanation for M1 activation. This means regions surrounding the M1 were

activated during mental rotation, and the strong interconnection between these regions caused

activity to present.

Bode et al. (2007) also centred their study around internal and external mental rotation

strategies. Participants were subject to a standard mental rotation task with stimuli intended to

trigger the use of an internal (e.g. hands) or external (e.g. abstract figures) strategy. Like

Eisenegger et al. (2007), single-pulse TMS was used to quantify M1 activity levels and measure

MEPs. Contrary to their hypothesis, there was no difference between the two stimulus

categories, and they concluded that M1 activation was not strategy-dependent but had a direct

involvement and potentially a spill-over effect. Despite employing the experimental design from

Eisenegger et al. (2007) and predicting results similar to that of Kosslyn et al. (1998), Bode and

colleagues produced results that contradicted both studies to varying degrees.

Previous studies have explored and confirmed a correlation between the M1 and mental

rotation. However, these findings contradict each other and generally lack exploration into a

causal relationship. This study aimed to investigate the presence of a causal relationship
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between the M1 and the mental rotation of Shepard Figures and hands, using single pulse

(‘neural noise’) transcranial magnetic stimulation (TMS). A one-tailed paired samples t-test

compared the RTs for real and sham TMS, an increase in RTs for real TMS would imply a

causal relationship. It is hypothesised that the M1 has a causal involvement in the mental

rotation of abstract objects. Therefore, it is predicted that participants who undertake the mental

rotation of Shepard figures will produce increased reaction times (be slower) under real TMS

conditions compared to sham TMS conditions. It is further hypothesised that the M1 has a

causal involvement in the mental rotation of hands. Hence, participants will have increased

reaction times when experiencing real TMS compared to sham TMS.

Method

Participants

Sixty participants took part in the study, 30 of whom were allocated to the hands

condition (16 female, 14 male; aged 18-30 years; M= 24.03, SD= 3.33) and 30 of whom were

allocated to the Shepard figures condition (15 female, 15 male, aged 18-29 years; M = 23.60,

SD = 2.87). All participants were right-handed, had normal or corrected-to-normal vision, and

had no history of psychological or neurological disorders.

Materials and Measures

A transcranial magnetic stimulator (TMS) equipped with a figure eight coil was used.

This was placed tangentially over the hand area of the left primary motor cortex with the handle

pointing backward and rotated 45 away from the midline. Stimuli were presented on the 21” flat

screen of a Dell PC using the Presentation Software Package. Stimuli were either Shepard

figures (Shepard & Metzler, 1971) or hands (Bode et al., 2007) which were approximately

constant in size (maximum of 10cm).

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Procedure and Design

Participants were seated approximately 70 cm (+/- 4) from the computer screen. Their

task was to mentally rotate the left object to see if it could be made to match the object on the

right. A positive match could be made in half of the trials, whereas in the other half the object

was mirrored. The degree of rotation necessary to come to either a “same” or “different”

judgement ranged between 45 and 315 degrees. For each category there were three objects,

each presented at four different viewing angles. All stimuli were shown twice, once for “same”

and once for “different”, leading to 24 trials in total per block for each condition which were

randomized.

Participants were instructed to mentally rotate each object to decide whether it was the

same or different. They were asked to do this as quickly and accurately as possible indicating

their decision using one of two buttons with their left hand. There were six experimental blocks 5

of which were preceded by a practice block. On odd blocks real TMS was applied (i.e., a TMS

pulse was applied 400 ms after stimulus onset at a pulse strength which was 110% above each

participant’s motor threshold). On even blocks sham TMS was applied (i.e., TMS coil is not

discharged). An average mean RT for TMS blocks and Sham blocks, for both hands and

Shepard Figures conditions was obtained and compared using a one-tailed paired samples t-

test.
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Results

Reaction times, in milliseconds, for the mental rotation of Shepard Figures and Hands

under real TMS and sham TMS conditions were summed and averaged.

Figure 1

Mean Reaction Times (ms) for Shepard Figures and Hands under Real and Sham Transcranial

Magnetic Stimulation (TMS) Conditions

Note. Error bars represent standard errors.

A one-tailed paired samples t-test measured the mean difference between RTs for the

mental rotation of Shepard Figures and Hands under real and sham TMS conditions. The

mental rotation of Shepard Figures under real and sham TMS conditions produced a mean

difference of 164.91 ms, with a standard error difference of 42.96. The difference in RTs would

be considered statistically significant (t(29)=3.84, p<0.001, d=0.70), and Cohen’s value d=0.70

suggests a medium effect size. The mean difference for mental rotation of hands under the two

conditions was 125.17 ms, with a standard error difference of 37.95. Similarly, these results

were significant (t(29)=3.30, p=0.0013, d=0.60), and the size effect was also medium.
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Discussion

This study aimed to investigate a causal relationship between the primary motor cortex

(M1) and the mental rotation of Shepard Figures and hands. Both hypotheses were supported

as there was an increase in RT for real TMS compared to sham in both experimental conditions,

with Shepard Figures producing higher results.

The findings by Kosslyn et al. (1998) both endorse and contradict this study. They found

that the M1 was active during the mental rotation of hands but not Shepard Figures because of

internal and external mechanisms, respectively. Internal mechanisms may explain the increase

in RT for the hands condition, however, Kosslyn et al. (1998) saw no evidence of M1 activity in

the Shepard Figures condition due to external mechanisms. This conclusion does not mirror the

current findings where the larger RT was for Shepard Figures. This misalignment may have

occurred as TMS has a higher spatial and temporal resolution than PET (Garcia-Sanz et al.,

2022). Resembling Eisenegger and colleagues (2007), the current findings found strong M1

activation in the mental rotation of Shepard Figures. Nevertheless, the procedure of the 2007

study likely influenced participants to use an ‘internal mechanism’, potentially swaying the

results to align with theories similar to Kosslyn et al. (1998). Eisenegger et al. (2007) contributed

their findings to the spill-over effect, and this study supported a direct involvement of the M1 in

mental rotation, differentiating the two studies as correlational and causal. Bode et al. (2007)

strongly reflects the current findings as they compared the M1 activation for Shepard Figures

and hands stimuli. They found no difference between the two and determined that M1 activity

was directly involved with mental rotation. There was a minimal difference between conditions in

this study, also supporting a direct M1 effect. Bode et al. (2007) relied on the measurement of

MEPs to form their conclusions and this study relied on RTs. Whilst similar results were

achieved, it is not possible to confirm the validity and reliability of these similarities given the

divergent procedures.
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For future applications, a larger, more diverse sample would increase reliability and

further investigate cognitive differences at an individual level. It would also be beneficial to

diversify the rotation stimuli to consolidate the correlational or causational role of the M1,

specifically regarding the uncertainty around the strategy dependence of M1 activation. From

the studies mentioned, M1 involvement in mental rotation is clear, but inconclusive results are

prevalent. For that reason, proof of a causal involvement would require further investigation.

To conclude, the findings of this study suggest a causal relationship between the M1 and

mental rotation and support a multi-faceted mind. Previous studies also provide strong support

for varying correlational theories. TMS and causational relationships between brain regions and

particular tasks can aid in disease prevention, clinical applications and cognitive enhancements.
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