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Collège de France, Feb 1st 2011Stanislas Dehaene Seminar
The construction of an experience of our own bodyown body
Henrik EhrssonD t t f N i K li k I tit t tDepartment of Neuroscience, Karolinska Institutet
Stockholm, Sweden
Body ownership y p
Whi h i h d?Which is my hand?
9 februari 2011Henrik Ehrsson 2
Asomatognosia andAsomatognosia and Somatoparaphrenia
Baier and Karnath (2008) St k
Arzy et al, . (2006) Arch Neurol
Stroke
B ti t l (2005) S i
9 februari 2011Henrik Ehrsson 3
Berti et al. (2005) Science
Body ownership as a’multisensory’ problemmultisensory problem
Whi h i h d?Which is my hand?
Henrik Ehrsson 9 februari 2011 4
OutlineOutline
1. Body ownership as a multisensory problemy p y p
2. Behavioural evidence from psychology
3. Multisensory responses in premotor and i t i t l h i l & i iintraparietal areas: neurophysiology & imaging
4 Imaging illusions of limb ownership4. Imaging illusions of limb ownership
5 Behavioural & imaging studies of full-body5. Behavioural & imaging studies of full-body ownership
6. Epilogue: applications to advanced prosthetics
1 Key principles of1. Key principles of multisensory integration
Henrik Ehrsson 9 februari 2011Stein and Stanford (2008) Nat Rev Neuroscience
Key principles ofKey principles of multisensory integration
• Temporal congruency principle.
• Spatial congruency principle.
• Principle of inverse effectiveness.
• Multisensory fusion problem.
M lti i t bl
• Multisensory assignment problem (causal interference problem)
H l d S (2005)
Henrik Ehrsson 9 februari 2011
Multisensory assignment problem From Körding et al. (2007)
Holmes and Spence (2005) Current Biolology
Body ownership as aBody ownership as a multisensory assignment problem
Seen &Felt hand Seen hand Felt hand
CVIS CTACCPRO CVIS CTAC
CPRO
Henrik Ehrsson 9 februari 2011
2. Behavioral evidence from experimental psychologyexperimental psychology
Henrik Ehrsson 9 februari 2011
Rubber hand illusion
9 februari 2011Henrik Ehrsson 10
Botvinick and Cohen (1998) Nature 391: 756
Measures of the illusionMeasures of the illusion
• Questionnaire with visual analogue scales• Questionnaire with visual analogue scales. (Botvinick and Cohen 1998)
• Hand perceived closer to the location of the rubber hand (proprioceptive drift): pointing error when asked to point to one’s hand (error when asked to point to one s hand (e.g Taskiris and Haggard 2005; Ehrsson et al. 2005)
• Emotional responses when the hand is ‘threatened’ or ‘hurt’ (e.g Armel & Ramachandran 2003; Ehrsson et al 2009)2003; Ehrsson et al. 2009)
• Visual capture of tactile localization (cross p (modal congruency task) (Pavani et al 2001; Zopf et al. 2009)
Behavioural evidence that the illusion depends on integration of visual, tactile and proprioceptive information in hand-centered reference frame
Temporal congruency principle:
• The brushstrokes applied to the two hands must be synchronous (temporal congruency vision-touch)(Botvinick & Cohen 1998; Armel & Ramachandrantouch)(Botvinick & Cohen 1998; Armel & Ramachandran2003; Ehrsson et al. 2004; Tsakiris and Haggard 2005)
L t l d l b k th ill i (>300• Longer temporal delays break the illusion (>300 ms) (temporal congruency vision-touch)(Shimada et al. 2009)
Behavioural evidence that the illusion depends on integration of visual, tactile and proprioceptive information in hand-centered reference frame
Spatial congruency principle:
• The brushstrokes must be applied to corresponding parts of the two hands (spatial congruency vision-touch)(e.g. Ehrsson et al. 2009)
• The rubber hand must be anatomically aligned with the hidden real hand. (spatial congruency vision-proprioception) (Pavani et al. 2001; Ehrsson et al. 2004; Tsakiris et al. 2005)
Tsakiris et al. (2005)
Behavioural evidence that the illusion depends on integration of visual, tactile and proprioceptive information in hand-centered reference frame
Hand-centered reference frames:
• The rubber hand is placed within reaching distance (30 cm from the real hand) (Ll d 2007)hand) (Llyod 2007).
• The direction of the brushstrokes must congruent in hand-centered reference frames, and not in allocentric external coordinates (Constantini and Haggard 2007)
Lloyd (2008)
3 N h i l3. Neurophysiology: multisensory integration in y gpremotor and intraparietal cortex
Henrik Ehrsson 9 februari 2011
Multisensory neurons in premotor cortex and peri personal spacecortex and peri-personal space
Henrik Ehrsson 9 februari 2011 16
From Rizzolatti et al. (1981) Exp. Brain Res.
Multisensory neurons andMultisensory neurons and peri-personal space
Fogassi et al. (1996) J. Neurophysiol.Hyvärinen & Poranen 1974Rizzolatti et al. 1981Graziano & Gross 1993 1997
Henrik Ehrsson 9 februari 2011 17
Graziano & Gross 1993, 1997 Duhamel et al. 1998
M lti dMultisensory neurons and body-part-centered coordinates
Henrik Ehrsson 9 februari 2011 18
From Graziano et al. (1997, 1999)
Multisensory responses in premotor cortex in anethesized monkeysy
PMv
Adopted from Graziano et al. 1999, 2000Graziano et al. (2000) Exp. Brain Res. Graziano et al. (2007) J. Neurophysiol.
M lti i t ti i t lMultisensory integration in ventral intraparietal area: super-additivity
Henrik Ehrsson 9 februari 2011 20Avillac et al (2007) Nature Neuroscience
Multisensory integration in ventral y gintraparietal area: spatial and temporal congruency effectscongruency effects
Temporal congruency Spatial congruency
Henrik Ehrsson 9 februari 2011 21
Avillac et al (2007) Nature Neuroscience
Functional magnetic resonance imaging (fMRI)imaging (fMRI)
BOLD
sig
nal
B
Ownership measure
Bl d d d t i l (BOLD)9 februari 2011Henrik Ehrsson 22
• Blood-oxygen-dependent signal (BOLD)
Human premotor and intraparietal cortices
Di t i f th i ht h d
Human premotor and intraparietal cortices performs multisensory integration
Direct view of the right hand:X
From participant’s perspective
Tactile stimulation Visual stimulationperspective
Human premotor and intraparietal p pcortices performs multisensory integrationg
Henrik Ehrsson
Gentile et al. (2010) J. Neurophysiol
Human premotor and intraparietalHuman premotor and intraparietal cortices performs multisensory integrationintegration
Henrik EhrssonGentile et al. (2010) J. Neurophysiol
fMRI evidence for peripersonal space in
3 sec 3 sec
p p phuman premotor and intraparietal cortices
3 sec
3 sec
Inter-trial3-11 sec
NearNear 3 sec
3 sec
FarNear
3 11 secFar Far
NEARFAR
100 cm
HAND
NEAR≈ 2 cm
100 cm
NO HAND
FIXATION50 cm
Henrik Ehrsson Brozzoli et al. (In manuscript)
fMRI-adaptation evidence for peripersonal space in human premotor and intraparietal cortices
Near
First
Second
Near
Far
Brozzoli et al. (In manuscript)
fMRI-adaptation evidence for i l i hperipersonal space in human
premotor and intraparietal cortices
Brozzoli et al. (In manuscript)
4. Imaging illusions of body ownershipownership
Henrik Ehrsson 9 februari 2011
fMRI f bb h d ill ifMRI of rubber hand illusion
Ehrsson et al. (2004) Science
Illusion ratingsIllusion ratings7
5
6at
ing
S C
4
5
bjec
tive
ra Sync CongrAsync CongrSync IncongrAsync Incongr
2
3Sub
1Feeling thatthe rubberhand is my
Touch feelingis located onrubber hand
Real handturningrubbery
Real hand isdrifting
towards the
It seems thatI have more
than oneyhand
yrubber hand right hand
Interaction brushstroke-timing and arm orientation (F(1, 17)=14.22; p<0.005)
Ehrsson et al. (2004) Science
Experimental paradigmp p g
Ehrsson et al. (2004) Science
Premotor activity reflects feeling ofPremotor activity reflects feeling of ownership (superadditative effect)
Left and right ventral premotor cortex (p<0.05 corrected)g p (p )
Ehrsson et al. Science 305:875-7. 2004Ehrsson et al. (2004) Science
Parietal activity reflects congruent hand y gposition and synchrony (additative effect)
Conjuction of main effects of synchrony andConjuction of main effects of synchrony and anatomical congruency (p<0.001 corrected)
Ehrsson et al. Science 305:875-7. 2004Ehrsson et al. (2004) Science
C l ti b t ill iCorrelation between illusion ratings and premotor activityg p y
Ehrsson et al. Science 305:875-7. 2004Ehrsson et al. (2004) Science
Conclusion: multisensory integration in premotor and intraparietal areasin premotor and intraparietal areas reflect limb ownership
Seen &Seen &Felt hand
CVIS C CPRO
Botvinick (2004) Science
CTACPRO
9 februari 2011Henrik Ehrsson 36
Somatic rubber hand illusion illusion
R. precentral gyrus (45 -6 57)
Premotor cortexPremotor cortex
0.1
0.5
0.9
ontr
aste
stim
ates
(a.u
.)
L. Inf precentral s. (-60 9 9)
0.9ates
-0.3 Illusion Async Incogr
Condition
Co
Intraparietal cortex
-0.3
0.1
0.5
0.9
Illusion Async IncogrCon
tras
test
ima
(a.u
.)
2 10 6 3 200
Condition
Ehrsson et al. J Neurosci. 25:10564-73, 20059 februari 2011Henrik Ehrsson 37
Ehrsson et al. J Neurosci. 25:10564-73, 2005
Re-calibration of position sense
Botvinick and Cohen (1998) Nature
9 februari 2011Henrik Ehrsson 38
Recalibration of position sense in aRecalibration of position sense in a ‘reaching circuit’ before illusion onset
Supplementary motor area, dorsal premotor cortex, th t i i t l t d th b llthe posterior parietal cortex, and the cerebellum
Henrik EhrssonEhrsson et al. (2004) Science
Objective evidence by th t i th bb h dthreatening the rubber hand
Ehrsson et al. (2007) PNAS 104:9828-9833 9 februari 2011Henrik Ehrsson 40
E i t l d iExperimental design
Eh t l (2007) PNAS 104 9828 9833Ehrsson et al. (2007) PNAS 104:9828-9833
9 februari 2011Henrik Ehrsson 41
I d th t k dIncreased threat-evoked response when owning the hand
Insula ACC
(p<0.05 corrected) (p<0.05 corrected)
9 februari 2011Henrik Ehrsson 42Ehrsson et al. (2007) PNAS 104:9828-9833
Activity in multisensory areas predict the threat-evoked response
ivity
(a.u
.) em
otor
acti
L. Insula activity (parameter estimates)
L. P
re
9 februari 2011Henrik Ehrsson
43
Ehrsson et al. (2007) PNAS
A preliminary model of limbA preliminary model of limb ownership
SMA, PMdHand-
S1 Processing of visual and tactile
signals in early sensory areas
PMvVisual areas
S1 Integration of visual and tactile
signals in posterior parietal cortex.
CB Recalibration of position sense in
motor regions.
Match of temporally and spatially congruent multisensory signals in premotor cortex (PMv) producepremotor cortex (PMv) produce ownership
Changes in other brain systems, e.g. in th ti l t
Henrik Ehrsson 9 februari 2011 44
the emotional system
Three-arm-illusion: k bl f t tia remarkable case of construction
Henrik Ehrsson 9 februari 2011 45
Ehrsson (2009) Perception 38, 310-312
Objective evidencej
Henrik Ehrsson 9 februari 2011 46
Ehrsson (2009) Perception 38, 310-312
Three-arm-illusion: k bl f t tia remarkable case of construction
50%50%
Seen &Felt hand
Seen &Felt hand
CVIS CTACCPRO
9 februari 2011
Henrik Ehrsson 9 februari 2011 47
Ehrsson (2009) Perception 38, 310-312
5. Extending the model to ownership of entire bodiesownership of entire bodies
O t f b d i (F B 2002)
9 februari 2011Henrik Ehrsson 48
Out-of-body experiences (From Brugger 2002)
Neurological out-of-body experiencesNeurological out of body experiences
Illusory body
Blanke et al. (2000) Nature Physical body
9 februari 2011Henrik Ehrsson 49
Blanke et al. (2002) Nature; Blanke and Mohr (2005) Brain Res. Reviews
First-person perspective and body ownershipbody ownership
Gibbson (1972) The ecological approach to visual perception (1972)
Ernst Mach (1885) Analyse der Empfindungento visual perception (1972)
9 februari 2011Henrik Ehrsson 50
Analyse der Empfindungen
Out-of-body illusion
9 februari 2011Henrik Ehrsson 51
Ehrsson (2007) Science 317: 1048
Experimental evidence
BA
Registration of skin conductance response (SCR)
9 februari 2011Henrik Ehrsson 52
Ehrsson (2007) Science
Ownership of an entire bodyOwnership of an entire body other than oneself
P tk & Eh t l PL S O (2008)
9 februari 2011Henrik Ehrsson 53
Petkova & Ehrsson et al. PLoS One (2008)
Obj ti t t Ski C d t RObjective test: Skin Conductance Responseo-
oho) Knife (Sync) Spoon (Sync) Knife (Async)
SCR
(mic
ro
Time (s)
Generalization of ownership to non stimulated body partsto non-stimulated body parts
1. Brush hand 2. Cut BellyKey experimental test:
SyncHand
AsyncHand
SyncBelly
AsyncBelly
559 februari 2011Henrik Ehrsson
Petkova & Ehrsson (2008) PLoS ONE
Effect of humanoid body shape
S A S ASyncBody
AsyncBody
SyncWood
AsyncWood
Petkova & Ehrsson PLoS One (2008)
Effect of visual perspective
First-person-tiperspective
(1PP)
Third-person-perspective
(3PP)
1PPSync
1PPAsync
3PPSync
3PPAsync
9 februari 2011Henrik Ehrsson 57
Petkova et al. In revision Frontiers in Cogntion
The illusion can be induced without video-technology, but only from the first-person perspective
1PP 3PP
p p p
Khoshnevis, Petkova & Ehrsson, in preparation, 2009Petkova et al. In revision Frontiers in Cognition
The illusion can be induced without video-technology, but only from the first-person perspective p p p
1PP SCR
3PP
IPPSync
IPPAsync
3PPSync
3PPAsync
Khoshnevis, Petkova & Ehrsson, in preparation, 2009
Sync Async Sync Async
Petkova et al. In revision Frontiers in Cognition
Imaging the full-body illusion:Imaging the full body illusion: Object x Timing
9 februari 2011Henrik Ehrsson 60Petkova et al. (submitted)
Interaction effect (Body X Sync)Interaction effect (Body X Sync)
I i h f ll b d ill iImaging the full-body illusion:Perspective x Timingg
Sync1-PP
Async1-PP
hird
PP
Sync AsyncPP
T
h
3-PP 3-PP
Firs
t
9 februari 2011Henrik Ehrsson
62
Petkova et al. (submitted)
Interaction effect (1PP X Sync)Interaction effect (1PP X Sync)
C l ti b t ill i t th dCorrelation between illusion strength and interaction BOLD effect
Imaging the full-body illusion: G li ti f hiGeneralisation of ownership across body parts
Timing
Sync1 PPne
quin
AsyncB ll
SyncB ll
Timing
AyncB ll1-PP
rt on
man BellyBelly
Manne
Belly
Bod
y pa Sync
1-PPSyncHand
AsyncHand
equin’s ha
Sync1-PP
SyncDetachedHand
AsyncDetachedHand
nd vs detaHand Hand ached han
Henrik Ehrsson
nd
Petkova et al. (submitted)
B d t ifi d b d t lBody-part specific and body-part general activations
Body-part specific and body-part generalBody-part specific and body-part general activations
Graziano et al. (2000) Exp Brain ResPetkova et al. (submittted)
Hand attached to mannequin Vs detachedHand attached to mannequin Vs detached hand (interaction effect)
Petkova et al. (submitted)
Generalization of ownership between body-specifc sections
Interaction (manniquin’s hand / detatched hand X Timing)
I t i l iInterim conclusion: ownerhship of entire bodies
Visual information from the first-person perspective, in conjunction with spatially and temporally congruent tactile, proprioceptiveand visual signals in ego-centric reference frames are necessary and sufficient factors elicit a sense of ownership of an entire body.
Our fMRI results suggest that the key neural mechanism is the integration of visual and somatic signals in body-part-centred
f f b lti l l ti ireference frames by multisensory neuronal populations in premotor and intraparietal cortex.
Importantly, these multisensory mechanisms operate in parallelfor different body parts with functional interplay between theseprocessesprocesses.
9 februari 2011Henrik Ehrsson 70
6. Applications: simulated bodies and advanced prostheticsand advanced prosthetics
Henrik Ehrsson 9 februari 2011
Projecting ownership to virtual bodies
Slater et al. (PLOS ONE 2010)( ) Jesper Mortensen, Pankaj Khanna, Insu Yu,
Bernhard Spanlang, Mel Slater EPSRC 'Presence in the Virtual Light Field' and FET PRESENCCIA.
Henrik Ehrsson 9 februari 2011 72
Humanoid robotic hand prostheses
SMART hand project (6FP project coordinated from University of Lund; Fredrik Sebelius Dept. Electrical Measurements, Lund University)Fredrik Sebelius Dept. Electrical Measurements, Lund University)
“the PISA hand” - developed and provided by the ARTS Lab, Scuola Superiore Sant´Anna (IST-2001-35094)
Henrik Ehrsson 9 februari 2011 73
p ( )
Upper limb amputees can be induced to e perience a r bber hand as their o nexperience a rubber hand as their own
Ehrsson HH Rosén B Stockselius A Ragnö C Rosen et al (2009) Scand J
Henrik Ehrsson 9 februari 2011 74
Ehrsson HH, Rosén B, Stockselius A, Ragnö C, Köhler P, Lundborg G (2008) Brain 131, 3443-3452
Rosen et al. (2009) Scand J Plast Reconstr Surg Hand
Surg. 43:260-266
Upper limb amputees can be induced to experience a rubber hand as their ownexperience a rubber hand as their own
Henrik Ehrsson 9 februari 2011 75Ehrsson et al. (2008) Brain 131, 3443-3452
SummarySummary• The problem of body ownership can be explained within
a multisensory framework
• The natural constraints of the rubber hand illusion and
a multisensory framework.
• The natural constraints of the rubber hand illusion and full-body illusions obey multisensory integration principles.
• Activity in premotor and intraparietal cortex obeys the ‘multisensory rules’ of the body ownership illusions; thusmultisensory rules of the body ownership illusions; thus multisensory integration in body-part-centered coordinates is a likely mechanism of ownership.
• By employing the multisensory principles of body ownership one can create a new type of arm prosthesis
9 februari 2011Henrik Ehrsson 76
ownership one can create a new type of arm-prosthesis that feels more like a real limb.
Brain, Body and Self Laboratory Dr. Alexander Skoglund, research
engineer
Dr. Hiske van Duinen, postdoc
Dr. Claudio Brozzoli, postdoc
Dr. Loretxu Bergouignan, postdoc
Dr Laura Schmalzl postdocDr. Laura Schmalzl, postdoc
Valeria Petkova, PhD student.
Giovanni Gentile, PhD student
Andreas Kalckert, PhD student.
htt // ki / h /
,
Arvid Guterstam PhD student
9 februari 2011
http://www.neuro.ki.se/ehrsson/Björn van der Hoort, Hedvig Zetterberg and Chris Berger (master students) Postdoc positions open!
Thank you!
http://www neuro ki se/ehrsson/http://www.neuro.ki.se/ehrsson/Henrik Ehrsson 9 februari 2011 78
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