Vi ual body con inui y du ing
ac ion obse a ion a ec s mo o
co ical exci abili y
Ma a Ma amala-Gomez1,2,3,9, F ancesca F isco4,5,9, Giacomo Guidali4, Ca lo a Lega5,6,
Alejand o Beacco7, Nadia Bolognini4,5,8 & Angelo Ma a i a4,5
Body owne ship, he sense ha he body belongs o onesel , can be al e ed by inducing body
manipula ions in Vi ual Reali y, such as by inc easing he isual discon inui y be ween he a a a ’s
hand and body. Body ep esen a ion manipula ions can also in luence mo o co ex exci abili y. We
hypo hesized ha he deg ee o body con inui y be ween one’s body and he obse ed i ual body
would a ec owne ship eelings and impac mo o co ex exci abili y du ing ac ion obse a ion.
Pa icipan s obse ed i ual hand mo emen s om a i s -pe son pe spec i e wi h he i ual hand
p esen ed wi h di e en le el o connec ion wi h hei eal hand; he i ual hand could be pa o a
ull i ual body co-loca ed wi h he eal body (Full-Body condi ion), i could appea as connec ed o
an uppe limb isually discon inuous om he eal body (Uppe Limb condi ion), o he i ual hand
appea ed in isola ion, ully discon inuous (De ached Hand condi ion). Resul s showed inc eased
co icospinal exci abili y when body con inui y is highe (Full-Body and Uppe Limb). This e ec
was media ed by owne ship and disowne ship eelings, suppo ing he ela ionship be ween body
pe cep ion and mo o sys em.
Keywo ds Vi ual embodimen , Ac ion obse a ion, Non-in asi e b ain s imula ion, Mo o exci abili y
The sense o embodimen (SoE) has been de ined as he sense o ha ing a body, and he body can be conside ed
bo h he subjec and objec o medical science and p ac ice1. The capabili y o ou b ain o ha e a ep esen a ion o
ou body esul s in a men al cons uc ion composed o pe cep ions and ideas abou he dynamic o ganiza ion o
ou own body, in ol ing ision, ouch, p op iocep ion, in e ocep ion, mo o con ol, and es ibula sensa ions2.
Indeed, he SoE e e s o he sense o ha ing a body, achie ed h ough p ocessing he di e en senso y inpu s
a i ing a ou body, which a e in eg a ed and in e p e ed by ou b ain o c ea e a cohe en ep esen a ion o
ou sel es. Acco dingly, De Vignemon , de ines he SoE as “(an objec ) E is embodied i and only i some p ope ies
o E a e p ocessed in he same way as he p ope ies o one’s own body”3. The SoE is induced when he ollowing
componen s a e p esen : (i) he Sense o Body Owne ship, which is he eeling ha a speci ic body pa belongs
o one’s body4; (ii) he Sense o Agency, which is he sense o in ending execu ing ac ions and he eeling o
con olling one’s body mo emen s5; and (iii) he Sense o Co-loca ion, which is he one’s spa ial expe ience o
being inside a body6.
Imme si e i ual eali y (IVR) sys ems ha e been widely used o induce he SoE, showing how and o
wha ex en we can expe ience a i ual body as ou own7. Rema kably, e en simply obse ing a i ual body,
posi ioned spa ially aligned wi h one’s physical body, can induce owne ship owa ds i 6. IVR sys ems p o ide
unique ad an ages, such as he abili y o easily manipula e speci ic ac o s associa ed wi h he embodied i ual
body in a con olled way ha would ha dly be possible in physical eali y8. In de ail, i is possible o manipula e
body ep esen a ion in e ms o body con inui y (i.e. con inui y o he connec ion be ween he body and i s
pa s) po en ially p omo ing a dissocia ion be ween one’s physical body and he i ual body obse ed om an
egocen ic isual pe spec i e9,10. Body con inui y e e s o he isual and s uc u al cohe ence be ween di e en
1Depa men o Cogni ion, De elopmen and Educa ional Psychology, Ins i u e o Neu osciences, Uni e si y o
Ba celona, Ba celona, Spain. 2Cogni ion and B ain Plas ici y Uni , Bell i ge Biomedical Resea ch Ins i u e [IDIBELL],
L’Hospi ale de Llob ega , Ba celona, Spain. 3Uni e si y School o Heal h and Spo (EUSES), Uni e si y o Gi ona,
Gi ona, Spain. 4Depa men o Psychology and Neu oMI, Uni e si y o Milano-Bicocca, Milano, I aly. 5Mind and
Beha io Technological Cen e , Uni e si y o Milano-Bicocca, Milan, I aly. 6Depa men o B ain and Beha io al
Sciences, Uni e si y o Pa ia, Pa ia, I aly. 7Depa men o Compu e Science, Uni e si a Poli ècnica de Ca alunya,
Ba celona, Spain. 8Labo a o y o Neu opsychology, Depa men o Neu o ehabili a ion Sciences, IRCCS Is i u o
Auxologico I aliano, 20122 Milan, I aly. 9Ma a Ma amala-Gomez and F ancesca F isco con ibu ed equally. email:
[email p o ec ed]; [email p o ec ed]
OPEN
Scien i ic Repo s | (2025) 15:13364 1
| h ps://doi.o g/10.1038/s41598-025-97695-9
www.na u e.com/scien i ic epo s
pa s o he body. He e, we use he e m ‘body con inui y’ o e e o he isual and s uc u al cohe ence be ween
di e en pa s o he body. In IVR, i is possible o manipula e body con inui y by p esen ing i ual body pa s
as ei he de ached om he i ual body o misaligned wi h he eal body. Fo ins ance, i has been shown ha
isual discon inui y be ween he i ual hand and a m signi ican ly dec eased pa icipan s’ sense o owne ship
o e he i ual hand9,10. Thus, when a i ual limb is a ached o he es o he i ual body, i p omo es a
s onge sense o owne ship o e he i ual body. Con e sely, when he i ual hand is comple ely de ached
om he i ual body, he sense o owne ship is educed.
Mo eo e , manipula ions o body ep esen a ion ha e an impac on mo o pe o mance11 and on he
ac i a ion o he p ima y mo o co ex. In his ega d, changing he mo phological cha ac e is ics o he a ec ed
i ual limb using IVR can imp o e mo o unc ions in pa ien s wi h s oke12,13. A ecen MRI s udy e ealed
ha obse ing i ual body mo emen s while being embodied in a i ual body lead o s onge ac i a ion o
he mo o and p emo o co ex14. Simila ly, mo o imp o emen s a e obse ing i ual body mo emen s had
been shown in pa ien s wi h dis al adius ac u e15 and in pa ien s wi h s oke16. Indeed, obse ing i ual body
mo emen s while being embodied in a i ual body has shown posi i e e ec s in he ac i a ion o he mo o
and p emo o co ex14, as well as on mo o eco e y in pa ien s wi h dis al adius ac u e15 and in pa ien s wi h
s oke16.
Some e idence showed a ela ionship be ween he ac i a ion o he mo o co ex and he SoE when obse ing
in olun a y hand mo emen s e oked by ansc anial magne ic s imula ion (TMS) o e he p ima y mo o co ex
(M1)17. In ac , i is well known ha co icospinal exci abili y is enhanced du ing he (passi e) obse a ion o
biological mo emen s ( he so-called ‘mo o esonance’ phenomenon18, and he magni ude o his acili a ion is
modula ed acco ding o he ea u es o he depic ed ac ions19,20. Hence, one may pos ula e ha obse ing hand
mo emen s om a i s -pe son pe spec i e while ully embodied in a i ual body inc eases mo o co ical
acili a ion du ing ac ion obse a ion. E en hough some e idence demons a es ha i ual body discon inui y
dec eases he sense o owne ship owa ds he i ual body9,10, o da e, none in es iga ed he ole o i ual body
ep esen a ion in e ms o body con inui y on M1 co ical exci abili y du ing ac ion obse a ion. Based on p io
e idence21–23, we hypo hesized ha g ea e i ual body con inui y would enhance he sense o owne ship and
inc ease mo o co ical exci abili y du ing ac ion obse a ion.
The p esen s udy aimed o in es iga e he impac o obse ing i ual hand mo emen s om a i s -pe son
pe spec i e on mo o co ex exci abili y while ully imme sed in a i ual en i onmen . Speci ically, he s udy
coun ed wi h di e en condi ions in e ms o isual con inui y o he mo ing i ual hand wi h he body o
he obse e : (i) Full Body, whe e he i ual hand is in eg a ed in o a ull body i s -pe son pe spec i e, (ii)
Uppe Limb, whe e he hand is in eg a ed in o a i ual uppe limb ha is isually discon inuous om he
pa icipan ’s body, (iii) Hand de ached, in which he i ual hand in isola ion and ully discon inuous wi h he
pa icipan ’s body (see Fig.1). In all condi ions, he bodily s imuli we e obse ed om a i s -pe son pe spec i e
in an imme si e i ual en i onmen . Du ing he ac ion obse a ion ask, pa icipan s obse ed a s a ic (s a ic
condi ion, wi h he i ual hand a es ) o (mo emen condi ion showing he index inge abduc ion) mo ing
le i ual hand. To measu e M1 co ical exci abili y, mo o e oked po en ials (MEPs) induced by single-pulse
TMS o e he igh M1 we e eco ded om he pa icipan ’s le hand du ing he ac ion obse a ion ask.
He e, conside ing p e ious indings using he same isual s imuli o mo emen s22,24,25, we expec ed ha
MEPs eco ded du ing mo emen obse a ion would be highe han he ones eco ded du ing he obse a ion o
s a ic s imuli, sugges ing he p esence o mo o esonance du ing ac ion obse a ion. Fu he mo e, co icospinal
exci abili y acili a ion du ing ac ion obse a ion is speci ic o he muscles in ol ed in he obse ed ac ion, a
leas o simple mo emen s like he one we adop ed18,20. Fo ins ance, an s udy by Guidali and colleagues22,
showed ha du ing he obse a ion o index inge abduc ion mo emen s ( ha is, he same ac ion depic ed in
he IVR asks o he p esen wo k) MEPs enhancemen was de ec able only om he i s do sal in e osseus (FDI)
muscle bu no om o he muscles no in ol ed in he iewed hand mo emen (i.e., abduc o digi i minimi –
ADM –22). Hence, we eco ded MEPs om bo h FDI and ADM muscles. Embodimen eelings we e assessed
h ough a ques ionnai e9,26. Consis en wi h p e ious s udies demons a ing an enhancemen in co icospinal
exci abili y by he obse a ion o simila hand mo emen s using non-imme si e pa adigms (i.e., mo o
esonance e ec )21–23, we expec ed o ind his e ec only o he muscle in ol ed in execu ing he obse ed
mo emen . Fu he mo e, we expec ed ha bo h mo o esonance and embodimen eelings would inc ease as
he isual con inui y be ween he eal and i ual body inc eased.
Resul s
The expe imen ollowed a 3(Condi ion) × 2(Type o Mo emen ) × 2(Muscle) ac o ial wi hin-subjec design
(see Fig.1). We conside ed MEPs ampli ude as an index o mo o co ex exci abili y and ques ionnai e sco es as
a measu e o embodimen sensa ions (see Table1 o ques ionnai e de ails). We employed a mixed-e ec s design
o analyze ou dependen a iables (MEPs), wi h a mul ile el linea eg ession es (mixed unc ion in S a a),
conside ing as ixed e ec s: body con inui y condi ion (Full Body, Uppe Limb, De ached Hand), ype o i ual
hand mo emen (s a ic o mo ing), and muscle (FDI o ADM). Signi ican e ec s we e u he explo ed wi h
Pos -hoc Sche e es . We used a mul ile el linea eg ession es (mixed unc ion in S a a) o iden i y di e ences
in he embodimen ques ionnai e sco es ac oss condi ions.
To u he explo e he mechanism h ough which body con inui y in luences co icospinal exci abili y, we
conduc ed a media ion analysis using a seemingly un ela ed eg ession (su eg unc ion in S a a). This analysis
examined whe he he sensa ion o embodimen , expe ienced du ing i ual body exposu e, media es he e ec
o body con inui y on co icospinal exci abili y. In all analyses, we included pa icipan s as a andom e ec
a iable o p ope ly accoun o in e -subjec a iabili y.
Scien i ic Repo s | (2025) 15:13364 2
| h ps://doi.o g/10.1038/s41598-025-97695-9
www.na u e.com/scien i ic epo s/
Di e en deg ees o body con inui y be ween he eal and he i ual body modula e meps
ampli ude
The esul s showed a signi ican 3(Condi ion)×2(Type o Mo emen )×2(Muscle) in e ac ion (z=-2.67, p < 0.01,
see Table2 o u he de ails). Conside ing MEPs eco ded om he FDI muscle, we ound a signi ican inc ease
in he MEPs ampli ude in he Full Body (z = 5.38, p < 0.001) and Uppe Limb condi ions (z = 5.55, p < 0.001)
when obse ing hand mo emen s compa ed o he s a ic condi ion, hence showing a mo o acili a ion e ec in
hese condi ions (See Fig.2; Table3 o mul iple compa isons (She e es )). In con as , no signi ican di e ences
eme ged in he De ached Hand condi ion (z=-0.03, p > 0.9). The e o e, co icospinal exci abili y acili a ion by
Fig. 1. (A) Expe imen al Design. (i) Neu ona iga ional p ocedu es and es ing mo o h eshold ( MT)
de e mina ion. (ii) Single-pulse TMS o e he igh M1 du ing an Ac ion Obse a ion Task. Ac ion Obse a ion
Task in which a mo ing and a s a ic i ual hand we e displayed in an IVR en i onmen . Du ing he
IVR ask, he co ico-spinal exci abili y was assessed by applying single-pulse TMS o e he igh M1
while simul aneously eco ding MEPs om wo le -hand muscles (FDI and ADM). (iii) Ques ionnai es.
Adminis a ion o he Embodimen Ques ionnai e, Simula o Sickness Ques ionnai e, and Vi ual Reali y
Expe ience Ques ionnai e. (B) Expe imen al Condi ions. Pa icipan s we e exposed o h ee i ual
en i onmen condi ions ha di e ed o he isual con inui y o he hand-s imulus image o he obse e ’s
body. In he Full Body, pa icipan s we e ully imme sed in a 360° IVR en i onmen , obse ing he i ual
hand in eg a ed in a Full Body, i s -pe son pe spec i e. In he Uppe Limb, pa icipan s obse ed he i ual
hand in eg a ed in an Uppe Limb isually discon inuous om he pa icipan ’s body. In he De ached hand,
pa icipan s obse ed he i ual hand in isola ion, ully discon inuous wi h he pa icipan ’s body. (C)
Expe imen al Se -up. The igu e depic ed he pa icipan ’s pos u e and he expe imen al appa a us. MT es ing
Mo o h eshold, TMS ansc anial magne ic s imula ion, MEP mo o e oked po en ial, M1 p ima y mo o
co ex, IVR imme si e i ual eali y, ADM abduc o digi i minimi, FDI i s do sal in e osseus, HMD head
moun ed display.
Scien i ic Repo s | (2025) 15:13364 3
| h ps://doi.o g/10.1038/s41598-025-97695-9
www.na u e.com/scien i ic epo s/
Fig. 2. (A) Di e en deg ees o body con inui y be ween he eal and he i ual body modula e MEPs
ampli ude. The Full Body and Uppe Limb condi ions p esen ed highe MEPs in he FDI han he De ached
Hand condi ion in he S a ic condi ion. Du ing mo emen condi ions, he Full Body and Uppe Limb
condi ions p esen ed highe MEPs in FDI muscle compa ed o he De ached Hand condi ion. The Uppe Limb
condi ion p esen ed highe MEPs in he ADM, du ing mo emen condi ion, compa ed o he Full Body and
De ached Hand condi ion. (B) Mo ing i ual hand induced highe MEPs compa ed o s a ic i ual hand. The
Full Body and Uppe Limb condi ions p esen ed highe MEPs in he FDI in he mo ing i ual hand condi ion
compa ed o he s a ic i ual hand condi ion. The Uppe Limb condi ions p esen ed highe MEPs in he
mo ing i ual hand condi ion compa ed o he s a ic i ual hand in he ADM. In he boxplo s, he medians
a e shown as ho izon al lines, and he boxes a e he in e qua ile anges (IQR). The whiske s a e be ween max
(min sco e, lowe qua ile − 1.5 IQR) o min (max sco e, uppe qua ile + 1.5 IQR). I he e a e alues ou side
he whiske s, hese a e con en ionally called “ou lie s” and a e shown by (∘). *p < 0.05, **p < 0.01, ***p < 0.0001.
Coe . zpLowe , 95% CI Uppe , 95% CI
Condi ionUppe Limb −0 0.004 -0.10 0.917 − 0.076 0.068
Condi ionDe ached Hand −0 0.047 -1.28 0.200 − 0.119 0.025
Type o mo emen Mo emen 0.018 0.50 0.617 − 0.054 0.091
MuscleFDI 0.680 18.42 < 0.001 0.607 0.752
Condi ionUppe Limb×Type o mo emen Mo emen 0.075 1.44 0.151 − 0.027 0.176
Condi ionDe ached Hand×Type o mo emen Mo emen − 0.007 -0.14 0.889 − 0.109 0.095
Condi ionUppe Limb×MuscleFDI − 0.007 -0.14 0.885 − 0.110 0.095
Condi ionDe ached Hand×MuscleFDI − 0.157 -3.00 0.003 − 0.259 − 0.054
Type o mo emen 1×MuscleFDI 0.183 3.52 < 0.001 0.081 0.286
Condi ionUppe Limb×Type o mo emen 1×MuscleFDI −0 0.094 -1.27 0.204 − 0.238 0.051
Condi ionDe ached Hand×Type o mo emen Mo emen × MuscleFDI − 0.197 -2.67 0.007 − 0.341 − 0.052
Table 2. The able shows he esul s o he 3(condi ion)×2( ype o mo emen )×2(muscle) on meps ampli ude.
Signi ican alues a e in bold.
Owne ship Q1 Du ing he i ual eali y expe ience, I el as i he i ual en i onmen was eal
Q4 Du ing he expe imen al session, I el as I had mo e han one body
Agency Q2 Du ing he expe imen al session, I el like I had con ol o e he a m ep esen ed on he sc een.
Q6 Du ing he expe imen al session, I el ha e en i I mo ed my eal a m, he a m ep esen ed on he sc een would no mo e.
Disowne ship
Q3 Du ing he expe imen al session, i seemed like I was unable o mo e my eal hand.
Q5 Du ing he expe imen al session, i el like my eal hand was disappea ed.
Q7 Du ing he expe imen al session, I couldn’ ell whe e my eal hand was loca ed.
Table 1. Embodimen ques ionnai e s a emen s.
Scien i ic Repo s | (2025) 15:13364 4
| h ps://doi.o g/10.1038/s41598-025-97695-9
www.na u e.com/scien i ic epo s/
ac ion obse a ion eme ged in condi ions wi h highe isual body con inui y be ween he eal and he i ual
body (i.e., Full Body and Uppe Limb). Mo eo e , FDI MEP ampli udes we e signi ican ly highe du ing ac ion
obse a ion in he Full Body (z=-10.37, p < 0.001) and Uppe Limb (z=-9.65, p < 0.001) condi ions compa ed
o he De ached Hand condi ion. Addi ionally, he e was a signi ican di e ence in MEPs ampli ude in he FDI
muscle du ing he obse a ion o s a ic hand s imuli when compa ing he Full Body (z=-5.51, p < 0.001) and
Uppe Limb (z=-5.15, p < 0.001) condi ions wi h he De ached Hand condi ion (Fig.2; Table3 o m mul iple
compa isons (She e es )). These esul s sugges ha a highe deg ee o isual con inui y be ween he eal and
i ual body can enhance M1 ac i a ion du ing he obse a ion o a hand in bo h s a ic and dynamic condi ions.
A he same ime, MEPs ampli ude eco ded om he con ol muscle (i.e., ADM) did no signi ican ly
di e be ween he s a ic and mo ing condi ions in he Full Body (n.s) and De ached Hand (n.s). Unexpec edly,
MEPs acili a ion by ac ion obse a ion eme ged in he ADM muscle in he Uppe Limb condi ion (z = 4.67;
p < 0.001). Addi ionally, no signi ican di e ences we e obse ed in he MEPs ampli ude o he ADM muscle
ac oss condi ions du ing he obse a ion o s a ic hand s imuli (n.s). Howe e , signi ican di e ences we e ound
du ing ac ion obse a ion when compa ing he Full Body (z=-2.93, p < 0.01) and De ached Hand (z=-5.31,
p < 0.001) condi ions (see Fig.2; Table3 o mul iple compa isons (She e es )).
The esul s indica ed a g adien in he mo o acili a ion e ec ela i e o he deg ee o body con inui y.
Speci ically, in he Full Body condi ion, a muscle-speci ic esponse was obse ed, wi h a di e ence be ween
mo emen and s a ic condi ions eme ging exclusi ely in he FDI muscle. In con as , he Uppe Limb condi ion
p oduced a muscle-unspeci ic esponse, wi h he acili a ion e ec appea ing in bo h he ADM and FDI muscles.
No mo o acili a ion e ec was obse ed in he Hand De ached condi ion. See Table3 o de ailed able esul s.
Highe body con inui y be ween he eal and he i ual body induces a s onge sense o
embodimen owa d he i ual body
The epo ed le els o owne ship showed a s a is ically signi ican di e ence be ween he Uppe Limb (z=-81.48,
p < 0.001) and De ached Hand (z= -117.26, p < 0.001) condi ion compa ed wi h he Full Body condi ion, being
highe in he Full Body condi ion. Fu he , he e was a signi ican di e ence be ween he De ached hand and he
Uppe Limb condi ions (z = 35.86, p < 0.001), showing highe le els o owne ship in he uppe limb condi ion. In
addi ion, he e was a signi ican di e ence be ween condi ions in e ms o agency, whe e pa icipan s epo ed
highe sco es in he Full Body condi ion compa ed o he Uppe Limb (z= -55.19, p < 0.001), and De ached
Hand condi ion (z=-59.75, p < 0.001). Again, a signi ican di e ence be ween Uppe Limb and De ached Hand
condi ion was ound, showing highe sco e in he Uppe limb condi ion (z = 4.60, p < 0.001). O e all, pa icipan s
a ed signi ican ly highe sco es ela ed o disowne ship (Q3, Q5, and Q7) in he Full Body condi ion compa ed
o he Uppe Limb (z=--38.07, p < 0.001; z=-63.15, p < 0.001; z=-48.44, p < 0.001) and De ached Hand (z=-42.93,
p < 0.001; z=-73.05, p < 0.001; z=-55.04, p < 0.001) condi ions. Mo eo e , he esul s also showed highe sco es
ela ed o disowne ship in he Uppe Limb condi ion compa ed o he De ached Hand condi ion (z = 4.89,
p < 0.001; z = 9.96, p < 0.001; z = 6.65, p < 0.001) (Fig.3).
Media ion analyses, by using he seemingly un ela ed eg essions (su ) me hod which allows o analyze
mul iple eg ession equa ions when hei e o s (dis u bances) a e co ela ed27, p edic he possible ela ionship
be ween he sco ing epo ed in he i ual embodimen ques ionnai e and he MEPs esponses ob ained a e
being exposed o he h ee di e en condi ions in e ms o i ual body con inui y (Full Body, Uppe Limb, and
De ached Hand), also e ealed ha he sense o owne ship (Q1), and he disowne ship index (Q3, Q5, and Q7)
media es di e ences in MEPs ampli ude among he di e en i ual body con inui y condi ions. Resul s showed
a signi ican ela ionship be ween he independen ac o i ual body con inui y and he media ion a iable
sense o owne ship, and disowne ship (see Table4).
P esence o a eeling o imme sion du ing he VR expe ience
The assessmen o he sense o p esence (i.e., he belie o being in ano he wo ld han whe e he body is loca ed)
du ing he VR expe ience was conduc ed using he Vi ual Reali y Expe ience scale28,29 (see Supplemen a y
Table 1). O e all, pa icipan s seem o pe cei e hemsel es as being imme sed in a eal en i onmen , as i hey
we e isi ing i (M = 0.43; SE = 0.20). Speci ically conside ing he h ee dimensions, we obse ed a posi i e sco e
o each dimension (sense o being he e: M = 0.58; SE = 0.23; eeling eal o p esen in he i ual en i onmen :
M = 0.17, SE = 0.30; Locali y: M = 0.47; SE = 0.27). The collec ed sco es du ing he expe imen al sessions indica e
ha he pa icipan s el imme sed in he VR en i onmen and expe ienced a sense o p esence wi hin he i ual
en i onmen (see Supplemen a y Ma e ials o addi ional de ails).
Absence o side e ec s wi hin he imme si e i ual eali y en i onmen
The possible side e ec s o cybe sickness we e assessed a e he VR expe ience using he Simula o Sickness
Ques ionnai e30; Supplemen a y Table 2). This ques ionnai e comp ises 16 i ems designed o e alua e a ious
Mo emen S a ic
ADM FDI ADM FDI
Full body s. de ached hand n.s z=-9.01, p < 0.001 n.s z=-5.59, p < 0.001
Uppe limb s. de ached hand z=-5.31, p < 0.0001 z=-9.01, p < 0.001 n.s z=-4.89, p < 0.001
Uppe limb s. ull body z=-2.93, p < 0.01 n.s n.s n.s
Table 3. Mul iple compa isons She e es .
Scien i ic Repo s | (2025) 15:13364 5
| h ps://doi.o g/10.1038/s41598-025-97695-9
www.na u e.com/scien i ic epo s/
symp oms, u ilizing a ou -poin scale (0 = None, 3 = Se e e). By applying a subjec i e h eshold ha conside ed
pa icipan s epo ing only sligh symp oms (sco e = 1) o each i em, i was obse ed ha all pa icipan s
eco ded a aw sco e below 16 (M = 4.47; SE = 0.599). Ou esul s indica e ha side e ec s we e a ed, on a e age,
below he “sligh ” le el o all i ems. Impo an ly, no pa icipan s ound i necessa y o in e up he imme si e
expe ience due o he sensa ion o sickness. Mo e speci ically, we also conside ed he sco es ela ed o he wo
main ac o s o he ques ionnai e (i.e., Nausea and Oculumo o ac o s). A highe sco e was obse ed o he
Oculomo o ac o (M = 3.56, SE = 0.477) in compa ison o he Nausea Fac o (M = 0.906, SE = 0.231]. This inding
implies ha he side e ec s a ibu ed o ou VR en i onmen we e mo e ela ed o oculomo o symp oms, such
as eye s ain, di icul y on concen a ing and ocusing, blu ed ision, and he sensa ion o “Fullness o he Head,”
a igue, and headache. Mo e de ails a e epo ed in he Supplemen a y ma e ials.
Embodimen componen Coe icien Con idence in e al z p- alue
Owne ship (Q1) 0.16 0.12–0.19 9.63 < 0.0001
Agency (Q2) -0.004 -0.32–0.02 -0.33 0.74
Disowne ship index
(Q3, Q5, Q7) 0.57 0.32–0.81 4.50 < 0.0001
Table 4. Resul s o media ion analysis.
Fig. 3. Di e en deg ees o con inui y be ween he eal and he i ual body a ec sense o embodimen
owa d he i ual body. The e we e signi ican di e ences be ween condi ions o all he ques ions o he
i ual embodimen ques ionnai e, showing highe sco es o owne ship and agency in he Full Body condi ion
compa ed o he Uppe Limb, and De ached Hand condi ions. Fu he , he Uppe Limb condi ion p esen ed
signi ican highe sco es ela ed o owne ship and agency compa ed o he De ached Hand condi ion. In he
boxplo s, he medians a e shown as ho izon al lines, and he boxes a e he in e qua ile anges (IQR). The
whiske s a e be ween max (min sco e, lowe qua ile − 1.5 IQR) o min (max sco e, uppe qua ile + 1.5 IQR).
I he e a e alues ou side he whiske s hese a e con en ionally called “ou lie s” and a e shown by (∘). *p < 0.05,
**p < 0.01, ***p < 0.0001.
Scien i ic Repo s | (2025) 15:13364 6
| h ps://doi.o g/10.1038/s41598-025-97695-9
www.na u e.com/scien i ic epo s/
Discussion
The p esen s udy in es iga ed mo o co ex exci abili y modula ions while obse ing hand mo emen s unde
a ying condi ions o isual body con inui y be ween he i ual hand and he obse e ’s body. Pa icipan s we e
imme sed in a i ual en i onmen h ough an HMD and obse ed a i ual body om a i s -pe son in h ee
di e en condi ions o isual con inui y wi h he eal body: (i) Full Body, (ii) Uppe Limb, (iii) De ached Hand.
Body con inui y e e s o he isual and s uc u al cohe ence be ween di e en pa s o he body. In IVR, i is
possible o manipula e body con inui y by p esen ing i ual body pa s as ei he de ached om he i ual body
o misaligned wi h he eal body. In de ail, he esul s o he p esen s udy showed ha pa icipan s epo ed
highe eelings o embodimen in he Full Body and Uppe Limb condi ions (condi ions showing highe isual
con inui y be ween he eal and he i ual body), compa ed o he De ached hand condi ions (lowe body
con inui y). To measu e M1 co icospinal exci abili y, le -hand MEPs we e eco ded ollowing single-pulse
TMS applied o he igh M1 while pa icipan s obse ed a s a ic o mo ing le i ual hand. Embodimen
eelings we e assessed h ough a ques ionnai e. Resul s showed ha eelings o embodimen we e enhanced as
he isual con inui y be ween he i ual and eal body inc eased (i.e., Full Body, and Uppe Limb). Mo eo e ,
he Full-Body and Uppe Limb condi ions induced highe MEPs ampli ude in he FDI muscle ei he when he
i ual hand was s a ic o mo ing compa ed o he Hand De ached condi ions, sugges ing highe M1 mo o
exci abili y, ega dless o ac ion obse a ion. C ucially, hese wo condi ions induced highe MEPs on mo ing
han s a ic hand obse a ion ials, sugges ing, besides a mo e gene al acili a ion e ec on M1 exci abili y in
hese condi ions, also an inc ease o mo o esonance e ec s du ing ac ion obse a ion21,22. Finally, he esul s
indica e ha he e ec on M1 exci abili y is media ed by he subjec i e sense o owne ship and disowne ship, as
assessed h ough he ques ionnai e. Speci ically, media ion analyses e ealed ha di e ences in MEPs ampli ude
ac oss i ual body con inui y condi ions we e signi ican ly in luenced by he sense o owne ship owa d he
i ual body and disowne ship owa d one’s body. Ou indings unde sco e he cen al ole o hese sensa ions in
shaping mo o exci abili y, as media o s o he ela ionship be ween i ual body con inui y and MEP ampli ude.
The sense o owne ship owa ds a i ual body may be a o ed by body con inui y. When we look down a
ou own bodies, we know ha he limbs we see belong o us because hey main ain a con inuous connec ion wi h
ou shoulde s o hips. This ana omical connec ion be ween con iguous body pa s c ea es a sense o cohe ence
and belonging, ein o cing ou pe cep ion o he body as an in eg a ed whole. Indeed, con inui y be ween body
pa s is c ucially s o ed in he so-called body s uc u al desc ip ion, which is c i ical o body ep esen a ion
and may be logically linked o “con inui y”31. Acco ding o his, p e ious s udies demons a ed ha in VR, he
physical con inui y o he i ual hand wi h he shoulde is a c ucial ac o in inducing he subjec i e illusion
o owne ship o e a i ual a m9. Tie i and colleagues demons a e ha small isual incong uences in a body’s
isual in eg i y can undamen ally a ec he sense o body owne ship in VR10. Fu he mo e, he au ho s ound
ha passi e obse a ion o a i ual hand de ached om he body simila ly modula es he sense o owne ship
and agency in bo h s a ic and dynamic con ex s o he i ual hand obse ed om a i s -pe son pe spec i e.
Howe e , a ull sense o body con inui y be ween he eal and he i ual body can be ully achie ed when using
imme si e i ual eali y. I is well-known ha h ough non-imme si e VR, i is no possible o induce ull
body-illusion2. Indeed, some s udies ha e demons a ed ha he co-loca ion o he eal body wi h he i ual
body when using imme si e i ual eali y sys ems is c ucial o induce he sense o owne ship owa d he i ual
body32. This is in line wi h p e ious s udies using he well-known ubbe hand illusion (RHI), in which he
concep o bodily con inui y was linked o he cons ain o ana omical plausibili y33. Speci ically, in he RHI,
he ake hand is placed in an ana omically cohe en posi ion and pa ially occluded by a clo h esembling an
a m, he eby main aining he sense o bodily con inui y. Such s uc u al cons ain s a e c ucial o inducing he
illusion. In his ega d, o he s udies ha e shown ha body owne ship illusions dec ease as pos u al incong uence
be ween he eal body and he ake one inc eases34,35.
Such an impac o body discon inui y on embodimen in e ms o a sense o owne ship and agency was
p e iously e ealed by Newpo and P es on whe e he au ho s showed ha de aching he ip o he inge
des oyed he unde lying owne ship o he emaining s ump as well as o he ip i sel , e en when he ip was
unde pa icipan s’ con ol36. Mo eo e , i has been shown ha he me e manipula ion o he isual appea ance
o a i ual limb could in luence he subjec i e eeling o owne ship and he physiological esponses (skin
conduc ance esponse, SCR) associa ed wi h a h ea ening s imulus app oaching he i ual hand37. In his s udy,
he au ho s exposed he pa icipan o ou di e en i ual a m condi ions in e ms o body con inui y om a
i s -pe son pe spec i e: (i) a i ual body wi h he igh hand and o ea m connec ed by a physiological w is
(Full-Limb), (ii) connec ed by a hin igid wi e, (iii) disconnec ed due o a missing w is , and (i ) disconnec ed
wi h a missing w is plus a plexiglass panel posi ioned be ween he hand and o ea m. The esul s e ealed ha
isual discon inui y be ween he i ual hand and o ea m a ec ed he physiological eac i i y associa ed wi h
a i ual h ea in e ms o SCR. SCR was s onge when he i ual hand was connec ed o he es o he body
(Full-Limb and Wi e condi ions) compa ed o o he condi ions whe e he i ual hand was disconnec ed om
he i ual body37. In he p esen wo k, we u he demons a ed ha he isual con inui y be ween one’s body
and he i ual body can a ec embodimen sensa ions. Speci ically, embodimen eelings we e inc emen ally
s onge as he isual con inui y be ween he i ual and eal body inc eased. The p esen s udy esul s suppo ed
he ela ion be ween body ep esen a ion and he mo o sys em, highligh ing he impo ance o a cohe en isual
con inui y be ween he i ual and one’s own body in modula ing mo o exci abili y.
The pa e ns ound on co icospinal exci abili y highligh ha he muscle-speci ic MEP acili a ion du ing
ac ion obse a ion (i.e., highe MEPs o he muscle ac ually in ol ed in he obse ed mo emen 25 –FDI)
is ound only du ing he Full Body condi ion. In he Uppe Limb condi ion, M1 exci abili y du ing ac ion
obse a ion was enhanced o bo h FDI and ADM muscles compa ed o s a ic s imuli, e ealing a gene alized
co icospinal acili a ion a he sigh o he mo ing index inge . As was commen ed abo e (see in oduc ion
sec ion), muscle speci ici y is one o he main p ope ies cha ac e izing he mo o esonance phenomenon
Scien i ic Repo s | (2025) 15:13364 7
| h ps://doi.o g/10.1038/s41598-025-97695-9
www.na u e.com/scien i ic epo s/
o simple mo emen s18,38. O e all, i is highly possible ha he ea u es o he depic ed hand may ha e
con ibu ed o he pa e ns ound in he p esen wo k on co icospinal exci abili y19,20. Fo ins ance, he absence
o mo o esonance in he De ached Hand condi ion could be in e p e ed as e idence ha , in an imme si e
en i onmen , ac ion obse a ion pe se is no manda o y o acili a e co icospinal exci abili y. Ra he , he sense
o embodimen and isual con inui y owa d he uppe limb may be he key ac o s allowing he eme gence
o his phenomenon. Hence, in he Uppe Limb condi ion, FDI and ADM inc eased exci abili y canno be
in e p e ed as ‘mo o esonance’ gi en ha he muscle-speci ici y o he e ec is no de ec able. Con e sely, i
sugges s a gene al inc ease o M1 exci abili y du ing ac ion obse a ion, ega dless o he ype o iewed ac ion
(mo emen s o he index inge , bu no o he li le inge ), gi en ha he depic ed hand canno be ully el as
he pa icipan ’s own hand due o body discon inui y. S ill, his isual s imulus is mo e salien o isuomo o
mi o ing p ocesses and ac i a es he mo o sys em o a g ea e ex en ha he p e ious condi ion. Howe e ,
his ac i a ion is no ully supe imposable o he ypical, muscle-speci ic, mo o esonance phenomenon –
which ins ead is ound in he Full Body condi ion – likely due o a less specialized mo o sys em ec ui men
o compensa e o isual discon inui y. Hence, he complex pa e n ound on co icospinal exci abili y in he
di e en expe imen al condi ions may unde lie in e angled and complemen a y e ec s e lec ing mi o ing,
a en ional, and embodimen p ocesses. Likely, M1 mi o ec ui men in VR en i onmen ollows mo e
complex pa e ns a a iance wi h s anda d se ings whe e he mo emen is simply obse ed on a sc een in on
o he pa icipan s18,38. Howe e , om he p esen da a, ou sugges ion emains a specula ion. Fu he s udies
a e needed o cla i y be e he neu ophysiological subs a es unde lying ac ion obse a ion in imme si e
en i onmen s like he ones adop ed he e.
Rega dless o he speci ic modula ion esponsible o he MEP pa e ns, ou esul s u he sugges ha
s onge isual con inui y be ween he i ual and eal body (i.e., Full Body condi ion) is a i al condi ion
o inducing muscle-speci ic mo o esonance, a leas when he subjec is embodied in o a i ual body. This
e idence could be highly in o ma i e o u u e s udies and clinical ea men s exploi ing ac ion obse a ion VR
se ings39. I mus be no ed ha , in he p esen s udy, ega dless o he obse a ion o a s a ic o mo ing i ual
hand, pa icipan s p esen ed highe mo o co ex exci abili y in he FDI muscle du ing he Full Body and Uppe
Limb condi ions compa ed o he De ached Hand one. This e idence u he ad ises ha , in an imme si e
en i onmen and when he con inui y be ween he eal and i ual body is inc eased, e en simple obse a ion o
a s a ic hand would induce inc eased co ical ac i a ion in M1, as i he s imulus would be in e p e ed in e ms
o he po en ial mo emen o one’s body pa 40,41.
O e all, ou pa e ns sugges ha he sense o embodimen owa d he i ual body and he con inui y o
he i ual body wi h he eal one can ha e an impac on M1 exci abili y. Indeed, he esul s om his s udy
show ha he sense o owne ship and disowne ship owa d he eal body media ed he co icospinal exci abili y
e ec s. Acco dingly, p e ious in es iga ions demons a ed a ela ionship be ween he sense o embodimen and
mo o co ex ac i a ion when embodied in a i ual body15,42–44. In he s udy conduc ed by Gonzalez-F anco
and colleagues43, pa icipan s we e ully embodied in a i ual body obse ed om a i s -pe son pe spec i e.
The i ual igh hand was isually aligned wi h hei eal hand. E en - ela ed b ain po en ials (ERPs) we e
eco ded in wo condi ions: a condi ion whe e he pa icipan ’s i ual hand was a acked wi h a kni e and a
con ol condi ion whe e he kni e only s uck he i ual able. In his s udy, he au ho s obse ed mu- hy hm
E en -Rela ed Desynch oniza ion (ERD) and Readiness Po en ial nega i i y in he mo o co ex when he
i ual hand was h ea ened, as would be expec ed i he eal hand was h ea ened and he pa icipan ied o
a oid ha m43. Mu- hy hm desynch oniza ion has been highly ela ed o mo o co ex ac i a ion45. Mo eo e ,
ano he s udy has e ealed ha changing he physical cha ac e is ics o an embodied i ual body can inc ease
mo o co ex exci abili y44. In his s udy, he au ho s induced he “s one a m illusion” while pa icipan s we e
embodied in a i ual body and p o ided single-pulse TMS o e alua e changes in mo o co ical exci abili y
associa ed wi h he s one a m illusion. To in es iga e i he “s one illusion” a ec ed mo o con ol, pa icipan s
pe o med a eaching ask wi h he human and “s one” i ual body. The esul s indica ed ha he s eng h o
he illusion was associa ed wi h enhanced mo o co ical exci abili y measu ed by MEPs, sugges ing a so o
mo o compensa ion ha ing embodied he illuso y s i ness o he body. Mo eo e , i has been shown ha eeling
ully embodied in a i ual body can enhance mo o ehabili a ion p ocesses in pa ien s wi h uppe limb mo o
diso de s, along wi h a posi i e ela ionship be ween he unc ional eco e y o he uppe limb and he epo ed
le els o owne ship and agency o e he i ual a m15. Kil eni and co-au ho s also demons a ed ha inducing
he illusion o losing a i ual a m when obse ing he i ual body om a i s -pe son pe spec i e dec eases
MEPs o he ex enso digi o um communis muscle o he con ala e al senso imo o co ex46. This inding
sugges ed ha inducing sho - e m illuso y pe cep ion o missing a body-pa can igge inhibi o y e ec s on
co icospinal pa hways. Consis en wi h his s udy, ou esul s ha e also shown ha eeling disowne ship owa d
he eal body media es MEPs esponses. Addi ionally, p e ious wo k has p o ided physiological e idence o a
educ ion o M1 mo o exci abili y in he hand egion when he eeling o disembodimen owa d he eal hand
is induced by bodily illusions26.
The p esen s udy sheds new ligh on unde s anding he ole o i ual embodimen on mo o co ex
exci abili y. Resul s show how obse ing i ual hand mo emen s connec ed o he es o he i ual body o
i ual a ms om a i s -pe son pe spec i e inc eases mo o exci abili y in he con ala e al M1. Such an e ec is
mainly media ed by he sense o owne ship and disowne ship owa d he i ual body. These esul s may help in
designing new applica ions ha combine non-in asi e echnologies (IVR and TMS) o inc ease M1 exci abili y
in clinical popula ions p esen ing mo o diso de s.
Limi a ions o he s udy
This s udy has some limi a ions ha should be acknowledged. Fi s , o e all, he pa icipan s epo ed low sco es
ega ding he sense o embodimen in he ques ionnai e. E en hough he sco es ela ed o he sense o body
Scien i ic Repo s | (2025) 15:13364 8
| h ps://doi.o g/10.1038/s41598-025-97695-9
www.na u e.com/scien i ic epo s/
owne ship a e highe in he Full Body condi ion compa ed o he Uppe Limb and De ached Hand condi ions,
s ill he sco e a e abo e 1 in a Like scale going ange om − 3 o 3. One possible eason o his esul is ha he
sense o embodimen has been induced by using a 360º came a (body cap ion om a i s -pe son pe spec i e),
as in he s udy om13. E en hough he induc ion o sense o embodimen h ough 360º images has been
demons a ed, i is mo e complica ed o pe ec ly co-loca e he posi ion o he eal body wi h he i ual one.
Rega ding his, some s udies ha e shown ha co-loca ion is a c ucial aspec o induce he sense o embodimen
owa d he i ual body47. Gi en he demons a ed ole o mul isenso y co ela ions ( isuo- ac ile o isuo-
mo o co ela ions) in enhancing he sense o embodimen in i ual eali y compa ed o me e isual exposu e48,
u u e s udies using 360º ideo o induce he sense o embodimen will bene i om inco po a ing mul isenso y
co ela ions be o e he ac ion obse a ion ask o enhance he sense o embodimen .
Mo eo e , he sample size in his s udy was ela i ely small ega ding he media ion analyses, la ge sample
sizes a e necessa y o u u e esea ch o con i m and ex end hese esul s, ensu ing mo e obus and widely
applicable conclusions. Las ly, o ehabili a ion pu poses, i would be bene icial o es he e ec s using mo e
ecological mo emen s o inc ease he applicabili y o hese indings in a clinical se ing.
Me hods
Pa icipan s
Thi y- wo heal hy pa icipan s we e ec ui ed o he expe imen ( emale = 21, male = 11, mean
age ± SD = 23.2 ± 3.08, age ange = 19–29). All pa icipan s we e igh -handed as assessed by he Edinbu gh
Handedness In en o y36. All pa icipan s illed in a ques ionnai e di ec ed o e alua e any possible
con aindica ions o pa icipa e in TMS s udy (TMS Sa e y Ques ionnai e49). The s udy ecei ed e hical app o al
om he Uni e si y o Milano-Bicocca’s E hical Commi ee and conduc ed in acco dance wi h he s anda ds o
he Decla a ion o Helsinki. All pa icipan s p o ided w i en in o med consen o hei pa icipa ion in he
expe imen .
Expe imen al p ocedu e
Pa icipan s we e sea ed in a chai , wi h hei le a m es ing on he able posi ioned in on o hem and hei
igh a m es ing on he legs, unde he able, and ou o hei iew. This placemen eplica ed he posi ion o he
i ual body in he Full Body condi ion. Subsequen ly, he h ee phases o he expe imen began (Fig.1A): (i)
Neu ona iga ion p ocedu es and es ing Mo o Th eshold ( MT) de e mina ion. The expe imen al session began
wi h he localiza ion o he le -hand FDI ho spo and wi h he de e mina ion o he indi idual MT. (ii) Single-
pulse TMS o e he igh M1 du ing an Ac ion Obse a ion Task in an IVR scena io. Co icospinal exci abili y
was assessed by eco ding MEPs, elici ed by single-pulse TMS o e he igh M1, in bo h he le -hand FDI and
he ADM muscles while pa icipan s iewed s a ic o mo ing le - i ual hand images in a andomized o de
(i.e., Ac ion Obse a ion Task). Du ing he Ac ion Obse a ion Task, h ee di e en condi ions in e ms o he
isual con inui y o he hand-s imulus image o he obse e ’s body we e p esen ed in an IVR scena io om
a i s -pe son pe spec i e: Full Body, Uppe Limb, Hand De ached (Fig.1B). Du ing he Full Body condi ion,
pa icipan s obse ed he i ual hand in eg a ed in a ull-body pe spec i e, wi hou discon inui y be ween
he i ual and he pa icipan ’s body. Du ing he Uppe Limb condi ion, pa icipan s obse ed he i ual
hand in eg a ed in an Uppe Limb isually discon inuous om he pa icipan ’s body. In he De ached Hand
condi ion, pa icipan s obse ed he i ual hand in isola ion, ully discon inuous wi h he pa icipan ’s body.
The expe imen al condi ions we e coun e balanced among he pa icipan s.
The i ual eali y scena ios we e p esen ed h ough he HMD (Oculus Ques 2) connec ed o a compu e
(No ebook Dell Alienwa e; M15CPU: in el i7-8750H, 8GB RAM, GPU NVIDIA RTX 2070 Max-q, Fig.1C).
T ials andomiza ion and iming o he s imuli p esen a ion we e all implemen ed using Uni y 2021 so wa e.
(iii) Ques ionnai es. A he end o each condi ion, pa icipan s illed ou an adap ed ques ionnai e measu ing
he sense o Embodimen . Speci ically, he ques ionnai e assessed he sense o owne ship and agency owa d
he i ual a a a , and disowne ship o he eal body9,26. To accoun o he po en ial e ec s o i ual eali y,
we also adminis e ed wo addi ional ques ionnai es o measu e he le el o discom o o sickness expe ienced
(Simula o Sickness Ques ionnai e30), and he ex en o which pa icipan s el imme sed wi hin he i ual
en i onmen 28,29 (see Supplemen al Ma e ials, Tables1 and 2). The en i e expe imen ook app oxima ely 1h o
comple e.
VR ac ion obse a ion ask
Pa icipan s pe o med a s anda d Ac ion Obse a ion Task, which in ol ed he p esen a ion o wo pic u es
depic ing a le hand in ei he s a ic o mo ing condi ions (Fig.1A). Thus, wo dis inc ypes o isual s imuli
we e p esen ed: s a ic, whe e he hand emained s a iona y, and mo ing, in which he index inge pe o med an
abduc ion-adduc ion mo emen (adap ed om22,25). Pa icipan s obse ed and ocused on hese isual s imuli
p esen ed in h ee dis inc condi ions (Fig.1B), all in ol ing comple e imme sion in a VR en i onmen using
an HMD. (i) In he Full Body condi ion, pa icipan s we e imme sed in he expe imen al oom, which could
be explo ed om a 360-deg ee pe spec i e. In his se ing, he i ual body’s posi ion p ecisely ma ched ha o
he pa icipan ’s eal body, and he i ual en i onmen eplica ed he expe imen al labo a o y. This se up was
designed o gua an ee ha pa icipan s expe ienced he same iewpoin as i hey we e looking a hei own
bodies. To achie e his, we cap u ed 360-deg ee pho os (s e eoscopic images) wi h a 360° came a (Ins a360
ONE X2) moun ed on he o ehead o a sea ed emale o male expe imen e wi h hei a m es ing on he able.
This se ing ensu ed ha pa icipan s expe ienced he same pe spec i e o hei own pos u e and su oundings
and ha s imuli ma ched he gende o each pa icipan . (ii) In he Uppe Limb condi ion, a 180° pic u e o
he a m placed on he able agains a 360° black backg ound was p esen ed, i.e., simila o a cinema mode.
Impo an ly, pa icipan s in his condi ion did no see he i ual body: he e was a sepa a ion be ween he
Scien i ic Repo s | (2025) 15:13364 9
| h ps://doi.o g/10.1038/s41598-025-97695-9
www.na u e.com/scien i ic epo s/
