{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T22:45:05Z","timestamp":1790808305093,"version":"4.1.0"},"update-to":[{"DOI":"10.1371\/journal.pcbi.1008848","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2021,4,1]],"date-time":"2021-04-01T00:00:00Z","timestamp":1617235200000}}],"reference-count":53,"publisher":"Public Library of Science (PLoS)","issue":"3","license":[{"start":{"date-parts":[[2021,3,22]],"date-time":"2021-03-22T00:00:00Z","timestamp":1616371200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["IIS-1908115"],"award-info":[{"award-number":["IIS-1908115"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["NINDS R01NS105241"],"award-info":[{"award-number":["NINDS R01NS105241"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.ploscompbiol.org"],"crossmark-restriction":false},"short-container-title":["PLoS Comput Biol"],"abstract":"<jats:p>Our sense of touch helps us encounter the richness of our natural world. Across a myriad of contexts and repetitions, we have learned to deploy certain exploratory movements in order to elicit perceptual cues that are salient and efficient. The task of identifying optimal exploration strategies and somatosensory cues that underlie our softness perception remains relevant and incomplete. Leveraging psychophysical evaluations combined with computational finite element modeling of skin contact mechanics, we investigate an illusion phenomenon in exploring softness; where small-compliant and large-stiff spheres are indiscriminable. By modulating contact interactions at the finger pad, we find this elasticity-curvature illusion is observable in passive touch, when the finger is constrained to be stationary and only cutaneous responses from mechanosensitive afferents are perceptible. However, these spheres become readily discriminable when explored volitionally with musculoskeletal proprioception available. We subsequently exploit this phenomenon to dissociate relative contributions from cutaneous and proprioceptive signals in encoding our percept of material softness. Our findings shed light on how we volitionally explore soft objects, i.e., by controlling surface contact force to optimally elicit and integrate proprioceptive inputs amidst indiscriminable cutaneous contact cues. Moreover, in passive touch, e.g., for touch-enabled displays grounded to the finger, we find those spheres are discriminable when rates of change in cutaneous contact are varied between the stimuli, to supplant proprioceptive feedback.<\/jats:p>","DOI":"10.1371\/journal.pcbi.1008848","type":"journal-article","created":{"date-parts":[[2021,3,22]],"date-time":"2021-03-22T13:31:02Z","timestamp":1616419862000},"page":"e1008848","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":29,"title":["An elasticity-curvature illusion decouples cutaneous and proprioceptive cues in active exploration of soft objects"],"prefix":"10.1371","volume":"17","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-2774-2760","authenticated-orcid":true,"given":"Chang","family":"Xu","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8530-2352","authenticated-orcid":true,"given":"Yuxiang","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3137-3822","authenticated-orcid":true,"given":"Gregory J.","family":"Gerling","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2021,3,22]]},"reference":[{"key":"pcbi.1008848.ref001","doi-asserted-by":"crossref","first-page":"342","DOI":"10.1016\/0010-0285(87)90008-9","article-title":"Hand movements: A window into haptic object recognition","volume":"19","author":"SJ Lederman","year":"1987","journal-title":"Cogn Psychol"},{"key":"pcbi.1008848.ref002","doi-asserted-by":"crossref","DOI":"10.1093\/acprof:oso\/9780195173154.001.0001","volume-title":"Human Hand Function.","author":"LA Jones","year":"2006"},{"key":"pcbi.1008848.ref003","doi-asserted-by":"crossref","first-page":"4188","DOI":"10.1073\/pnas.1520866113","article-title":"Spatial patterns of cutaneous vibration during whole-hand haptic interactions","volume":"113","author":"Y Shao","year":"2016","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pcbi.1008848.ref004","doi-asserted-by":"crossref","first-page":"eaaw3121","DOI":"10.1126\/sciadv.aaw3121","article-title":"Touch as an auxiliary proprioceptive cue for movement control","volume":"5","author":"A Moscatelli","year":"2019","journal-title":"Sci Adv"},{"key":"pcbi.1008848.ref005","doi-asserted-by":"crossref","first-page":"4","DOI":"10.1109\/TOH.2019.2959767","article-title":"Tactile exploration strategies with natural compliant objects elicit virtual stiffness cues","volume":"13","author":"C Xu","journal-title":"IEEE Trans Haptics. 2020"},{"key":"pcbi.1008848.ref006","doi-asserted-by":"crossref","first-page":"2290","DOI":"10.1073\/pnas.1816278116","article-title":"Distinction of self-produced touch and social touch at cortical and spinal cord levels","volume":"116","author":"R Boehme","year":"2019","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pcbi.1008848.ref007","first-page":"437","volume-title":"IEEE World Haptics Conference (WHC).","author":"M Cavdan","year":"2019"},{"key":"pcbi.1008848.ref008","author":"U Neisser","year":"1976"},{"key":"pcbi.1008848.ref009","doi-asserted-by":"crossref","first-page":"273","DOI":"10.1109\/TOH.2011.2","article-title":"Tactile and haptic illusions","volume":"4","author":"SJ Lederman","year":"2011","journal-title":"IEEE Trans Haptics"},{"key":"pcbi.1008848.ref010","doi-asserted-by":"crossref","first-page":"742","DOI":"10.1016\/j.brainresbull.2008.01.008","article-title":"A brief taxonomy of tactile illusions and demonstrations that can be done in a hardware store","volume":"75","author":"V. Hayward","year":"2008","journal-title":"Brain Res Bull"},{"key":"pcbi.1008848.ref011","first-page":"122","article-title":"Analyse experimentale de quelques elements de la sensation de poids","volume":"3","author":"A. Charpentier","year":"1891","journal-title":"Arch Physiol Norm Pathol"},{"key":"pcbi.1008848.ref012","doi-asserted-by":"crossref","first-page":"e0236440","DOI":"10.1371\/journal.pone.0236440","article-title":"The size-weight illusion comes along with improved weight discrimination","volume":"15","author":"C Wolf","year":"2020","journal-title":"PLoS One"},{"key":"pcbi.1008848.ref013","doi-asserted-by":"crossref","first-page":"1002080","DOI":"10.1371\/journal.pcbi.1002080","article-title":"How haptic size sensations improve distance perception","volume":"7","author":"PW Battaglia","year":"2011","journal-title":"PLoS Comput Biol"},{"key":"pcbi.1008848.ref014","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1080\/00223980.1956.9713002","article-title":"A tactual-kinesthetic curvature illusion","volume":"42","author":"ES Edgington","year":"1956","journal-title":"J Psychol"},{"key":"pcbi.1008848.ref015","doi-asserted-by":"crossref","first-page":"1555","DOI":"10.1152\/jn.00719.2018","article-title":"Illusory changes in the perceived speed of motion derived from proprioception and touch","volume":"122","author":"A Moscatelli","year":"2019","journal-title":"J Neurophysiol"},{"key":"pcbi.1008848.ref016","doi-asserted-by":"crossref","first-page":"154","DOI":"10.1038\/349154a0","article-title":"A neurological dissociation between perceiving objects and grasping them","volume":"349","author":"MA Goodale","year":"1991","journal-title":"Nature"},{"key":"pcbi.1008848.ref017","doi-asserted-by":"crossref","first-page":"604","DOI":"10.1016\/S0960-9822(00)00132-9","article-title":"Separate neural pathways for the visual analysis of object shape in perception and prehension","volume":"4","author":"MA Goodale","year":"1994","journal-title":"Curr Biol"},{"key":"pcbi.1008848.ref018","doi-asserted-by":"crossref","first-page":"3887","DOI":"10.1152\/jn.00851.2005","article-title":"Opposite Perceptual and Sensorimotor Responses to a Size-Weight Illusion","volume":"95","author":"MS Grandy","year":"2006","journal-title":"J Neurophysiol"},{"key":"pcbi.1008848.ref019","doi-asserted-by":"crossref","first-page":"737","DOI":"10.1038\/76701","article-title":"Independence of perceptual and sensorimotor predictions in the size\u2013weight illusion","volume":"3","author":"JR Flanagan","year":"2000","journal-title":"Nat Neurosci"},{"key":"pcbi.1008848.ref020","doi-asserted-by":"crossref","first-page":"1159","DOI":"10.1016\/j.cub.2016.02.052","article-title":"The change in fingertip contact area as a novel proprioceptive cue","volume":"26","author":"A Moscatelli","year":"2016","journal-title":"Curr Biol"},{"key":"pcbi.1008848.ref021","doi-asserted-by":"crossref","first-page":"eaaw8845","DOI":"10.1126\/sciadv.aaw8845","article-title":"Role of indentation depth and contact area on human perception of softness for haptic interfaces","volume":"5","author":"C Dhong","year":"2019","journal-title":"Sci Adv"},{"key":"pcbi.1008848.ref022","doi-asserted-by":"crossref","first-page":"1535","DOI":"10.1177\/1541931218621347","article-title":"In the tactile discrimination of compliance, perceptual cues in addition to contact area are required","volume":"62","author":"C Xu","year":"2018","journal-title":"Proc Hum Factors Ergon Soc Annu Meet"},{"key":"pcbi.1008848.ref023","doi-asserted-by":"crossref","first-page":"10864","DOI":"10.1073\/pnas.1706233114","article-title":"Why pens have rubbery grips","volume":"114","author":"B Dzidek","year":"2017","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pcbi.1008848.ref024","first-page":"1906850","article-title":"Organic haptics: Intersection of materials chemistry and tactile perception","author":"DJ Lipomi","year":"2019","journal-title":"Adv Funct Mater"},{"key":"pcbi.1008848.ref025","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1109\/TOH.2009.16","article-title":"Cues for haptic perception of compliance","volume":"2","author":"WM Bergmann Tiest","year":"2009","journal-title":"IEEE Trans Haptics"},{"key":"pcbi.1008848.ref026","doi-asserted-by":"crossref","first-page":"232","DOI":"10.1109\/TOH.2017.2715845","article-title":"Force-rate cues reduce object deformation necessary to discriminate compliances harder than the skin","volume":"11","author":"SC Hauser","year":"2018","journal-title":"IEEE Trans Haptics"},{"key":"pcbi.1008848.ref027","doi-asserted-by":"crossref","first-page":"218","DOI":"10.1152\/jn.00624.2015","article-title":"Computational modeling indicates that surface pressure can be reliably conveyed to tactile receptors even amidst changes in skin mechanics","volume":"116","author":"Y Wang","year":"2016","journal-title":"J Neurophysiol"},{"key":"pcbi.1008848.ref028","doi-asserted-by":"crossref","first-page":"88","DOI":"10.1152\/jn.1995.73.1.88","article-title":"Tactual discrimination of softness","volume":"73","author":"MA Srinivasan","year":"1995","journal-title":"J Neurophysiol"},{"key":"pcbi.1008848.ref029","doi-asserted-by":"crossref","first-page":"242","DOI":"10.1109\/TOH.2011.19","article-title":"Exploratory strategies in haptic softness discrimination are tuned to achieve high levels of task performance","volume":"4","author":"L Kaim","year":"2011","journal-title":"IEEE Trans Haptics"},{"key":"pcbi.1008848.ref030","first-page":"353","volume-title":"IEEE World Haptics Conference (WHC)","author":"C Xu","year":"2019"},{"key":"pcbi.1008848.ref031","first-page":"1","article-title":"Visuo-Haptic Discrimination of Viscoelastic Materials","author":"O Caldiran","year":"2019","journal-title":"IEEE Trans Haptics"},{"key":"pcbi.1008848.ref032","doi-asserted-by":"crossref","first-page":"555","DOI":"10.1016\/j.neuron.2015.03.020","article-title":"Multimodal Interactions between Proprioceptive and Cutaneous Signals in Primary Somatosensory Cortex","volume":"86","author":"SS Kim","year":"2015","journal-title":"Neuron"},{"key":"pcbi.1008848.ref033","doi-asserted-by":"crossref","first-page":"170","DOI":"10.1038\/nn1177","article-title":"First spikes in ensembles of human tactile afferents code complex spatial fingertip events","volume":"7","author":"RS Johansson","year":"2004","journal-title":"Nat Neurosci"},{"key":"pcbi.1008848.ref034","doi-asserted-by":"crossref","first-page":"e1006264","DOI":"10.1371\/journal.pcbi.1006264","article-title":"Computation predicts rapidly adapting mechanotransduction currents cannot account for tactile encoding in Merkel cell-neurite complexes","volume":"14","author":"GJ Gerling","year":"2018","journal-title":"PLOS Comput Biol"},{"key":"pcbi.1008848.ref035","doi-asserted-by":"crossref","first-page":"360","DOI":"10.1007\/978-3-662-44196-1_44","volume-title":"Haptics: Neuroscience, Devices, Modeling, and Applications","author":"Y Wang","year":"2014"},{"key":"pcbi.1008848.ref036","doi-asserted-by":"crossref","first-page":"495","DOI":"10.3758\/BF03213075","article-title":"Manual discrimination of compliance using active pinch grasp: The roles of force and work cues","volume":"57","author":"HZ Tan","year":"1995","journal-title":"Percept Psychophys"},{"key":"pcbi.1008848.ref037","doi-asserted-by":"crossref","first-page":"1651","DOI":"10.1152\/physrev.00048.2011","article-title":"The proprioceptive senses: Their roles in signaling body shape, body position and movement, and muscle force","volume":"92","author":"U Proske","year":"2012","journal-title":"Physiol Rev"},{"key":"pcbi.1008848.ref038","doi-asserted-by":"crossref","first-page":"72","DOI":"10.1037\/0033-2909.103.1.72","article-title":"Motor Illusions: What Do They Reveal About Proprioception?","volume":"103","author":"LA Jones","year":"1988","journal-title":"Psychol Bull"},{"key":"pcbi.1008848.ref039","volume-title":"Detection theory: A user\u2019s guide","author":"NA Macmillan","year":"1991"},{"key":"pcbi.1008848.ref040","first-page":"1","article-title":"A Systematic Comparison of Perceptual Performance in Softness Discrimination with Different Fingers","author":"AC Zoeller","year":"2020","journal-title":"Attention, Perception, Psychophys"},{"key":"pcbi.1008848.ref041","doi-asserted-by":"crossref","first-page":"531","DOI":"10.3758\/BF03212187","article-title":"Manual discrimination of force using active finger motion","volume":"49","author":"XD Pang","year":"1991","journal-title":"Percept Psychophys"},{"key":"pcbi.1008848.ref042","doi-asserted-by":"crossref","first-page":"e1008081","DOI":"10.1371\/journal.pcbi.1008081","article-title":"Predicting precision grip grasp locations on three-dimensional objects","volume":"16","author":"LK Klein","year":"2020","journal-title":"PLOS Comput Biol."},{"key":"pcbi.1008848.ref043","doi-asserted-by":"crossref","first-page":"277","DOI":"10.1007\/BF00238156","article-title":"Factors influencing the force control during precision grip","volume":"53","author":"G Westling","year":"1984","journal-title":"Exp Brain Res"},{"key":"pcbi.1008848.ref044","first-page":"7","volume-title":"IEEE Haptics Symposium, HAPTICS","author":"SC Hauser","year":"2018"},{"key":"pcbi.1008848.ref045","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1113\/jphysiol.1995.sp020875","article-title":"Skin strain patterns provide kinaesthetic information to the human central nervous system","volume":"487","author":"BB Edin","year":"1995","journal-title":"J Physiol"},{"key":"pcbi.1008848.ref046","first-page":"72","volume-title":"Lecture Notes in Computer Science (including subseries Lecture Notes in Artificial Intelligence and Lecture Notes in Bioinformatics)","author":"A Moscatelli","year":"2014"},{"key":"pcbi.1008848.ref047","doi-asserted-by":"crossref","first-page":"20151661","DOI":"10.1098\/rspb.2015.1661","article-title":"The brain uses extrasomatic information to estimate limb displacement","volume":"282","author":"V. Terekhov A","year":"2015","journal-title":"Proc R Soc B Biol Sci"},{"key":"pcbi.1008848.ref048","volume-title":"Role of mechanics in tactile sensing of shape","author":"K. Dandekar","year":"1995"},{"key":"pcbi.1008848.ref049","first-page":"261","author":"RJ Gulati","year":"1995"},{"key":"pcbi.1008848.ref050","volume-title":"Psychophysics: The Fundamentals","author":"GA Gescheider","year":"1997","edition":"3"},{"key":"pcbi.1008848.ref051","doi-asserted-by":"crossref","first-page":"268","DOI":"10.1109\/TOH.2012.74","article-title":"Application of psychophysical techniques to haptic research","volume":"6","author":"LA Jones","year":"2013","journal-title":"IEEE Trans Haptics"},{"key":"pcbi.1008848.ref052","doi-asserted-by":"crossref","first-page":"451","DOI":"10.1109\/TOH.2019.2899298","article-title":"Effects of stimulus exploration length and time on the integration of information in haptic softness discrimination","volume":"12","author":"A Metzger","year":"2019","journal-title":"IEEE Trans Haptics"},{"key":"pcbi.1008848.ref053","first-page":"364","article-title":"Sequential response bias in psychophysical experiments","volume":"3","author":"WA Wagenaar","year":"1968","journal-title":"Percept Psychophys"}],"updated-by":[{"DOI":"10.1371\/journal.pcbi.1008848","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2021,4,1]],"date-time":"2021-04-01T00:00:00Z","timestamp":1617235200000}}],"container-title":["PLOS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1008848","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,4,1]],"date-time":"2021-04-01T14:47:11Z","timestamp":1617288431000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1008848"}},"subtitle":[],"editor":[{"given":"Adrian M.","family":"Haith","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2021,3,22]]},"references-count":53,"journal-issue":{"issue":"3","published-online":{"date-parts":[[2021,3,22]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1008848","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2020.08.04.237016","asserted-by":"object"}]},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,3,22]]}}}