{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,8]],"date-time":"2026-05-08T07:14:44Z","timestamp":1778224484500,"version":"3.51.4"},"reference-count":69,"publisher":"Springer Science and Business Media LLC","issue":"2","license":[{"start":{"date-parts":[[2026,2,28]],"date-time":"2026-02-28T00:00:00Z","timestamp":1772236800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,2,28]],"date-time":"2026-02-28T00:00:00Z","timestamp":1772236800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100005721","name":"Universit\u00e4t Bielefeld","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100005721","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Biol Cybern"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    Proprioception is key to all behaviours that involve the control of force, posture or movement. Computationally, many proprioceptive afferents share three features: First, their strictly local encoding of stimulus magnitudes causes range fractionation in sensory arrays. As a result, encoding of large joint angle ranges requires convergence of afferent information onto first-order interneurons. Second, their phasic-tonic response properties lead to fractional encoding of the fundamental sensory magnitude and its derivatives (e.g., joint angle and angular velocity). Third, the distribution of disjunct sensory arrays across the body implies that complex movements involve information from multiple joints or limbs. The present study proposes a multi-layer spiking neural network for distributed computation of whole-body posture and movement. The first part of the study models strictly local, phasic-tonic encoding of joint angle by proprioceptive hair field afferents by use of Adaptive Exponential Integrate-and-Fire neurons. Fractionally encoded afferent information about single-joint posture and movement converges on two types of first-order interneurons, tuned to encode either joint angle or velocity across the entire working range with high accuracy. In velocity-encoding interneurons, spike rate increases linearly with angular velocity. The companion paper exploits this distributed position\/velocity encoding in second- and third-order interneurons, using combinations of two or three position\/velocity inputs from disjunct arrays. The encoding properties of all interneuron layers are evaluated with experimental data on whole-body kinematics of unrestrained stick insect locomotion, comprising concurrent joint angle time courses of\n                    <jats:inline-formula>\n                      <jats:tex-math>$$6\\times 3$$<\/jats:tex-math>\n                    <\/jats:inline-formula>\n                    leg joints. The hierarchical model allows increasingly complex encoding of posture and movement, from angular velocity of a single joint, to movement cycle phases of an entire limb, to parameters of overall body posture.\n                  <\/jats:p>","DOI":"10.1007\/s00422-025-01032-2","type":"journal-article","created":{"date-parts":[[2026,2,28]],"date-time":"2026-02-28T08:17:45Z","timestamp":1772266665000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["A spiking neural network model for fractional proprioceptive encoding of limb posture and movement in insects"],"prefix":"10.1007","volume":"120","author":[{"given":"Thomas","family":"van der Veen","sequence":"first","affiliation":[]},{"given":"Yonathan","family":"Cohen","sequence":"additional","affiliation":[]},{"given":"Elisabetta","family":"Chicca","sequence":"additional","affiliation":[]},{"given":"Volker","family":"D\u00fcrr","sequence":"additional","affiliation":[]}],"member":"297","published-online":{"date-parts":[[2026,2,28]]},"reference":[{"issue":"9","key":"1032_CR1","doi-asserted-by":"publisher","first-page":"2099","DOI":"10.1152\/jn.00281.2013","volume":"110","author":"JM Ache","year":"2013","unstructured":"Ache JM, D\u00fcrr V (2013) Encoding of near-range spatial information by descending interneurons in the stick insect antennal mechanosensory pathway. J Neurophysiol 110(9):2099\u20132112","journal-title":"J Neurophysiol"},{"issue":"7","key":"1032_CR2","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pcbi.1004263","volume":"11","author":"JM Ache","year":"2015","unstructured":"Ache JM, D\u00fcrr V (2015) A computational model of a descending mechanosensory pathway involved in active tactile sensing. PLoS Comput Biol 11(7):e1004263","journal-title":"PLoS Comput Biol"},{"issue":"9","key":"1032_CR3","doi-asserted-by":"publisher","first-page":"4081","DOI":"10.1523\/JNEUROSCI.3350-14.2015","volume":"35","author":"JM Ache","year":"2015","unstructured":"Ache JM, Haupt SS, D\u00fcrr V (2015) A direct descending pathway informing locomotor networks about tactile sensor movement. J Neurosci 35(9):4081\u20134091","journal-title":"J Neurosci"},{"issue":"01","key":"1032_CR4","first-page":"217","volume":"1","author":"HB Barlow","year":"1961","unstructured":"Barlow HB et al (1961) Possible principles underlying the transformation of sensory messages. Sensory Commun 1(01):217\u2013233","journal-title":"Sensory Commun"},{"issue":"5","key":"1032_CR5","doi-asserted-by":"publisher","first-page":"661","DOI":"10.1007\/s00359-019-01355-z","volume":"205","author":"FG Barth","year":"2019","unstructured":"Barth FG (2019) Mechanics to pre-process information for the fine tuning of mechanoreceptors. J Comp Physiol A 205(5):661\u2013686","journal-title":"J Comp Physiol A"},{"key":"1032_CR6","unstructured":"Berg E (2014) Adaptive motor control: Neuronal mechanisms underlying (targeted) searching movements. Doctoral Dissertation, Universit\u00e4t zu K\u00f6ln, https:\/\/kups.ub.uni-koeln.de\/5773\/"},{"issue":"6","key":"1032_CR7","first-page":"1064","volume":"216","author":"E Berg","year":"2013","unstructured":"Berg E, B\u00fcschges A, Schmidt J (2013) Single perturbations cause sustained changes in searching behavior in stick insects. J Exp Biol 216(6):1064\u20131074","journal-title":"J Exp Biol"},{"issue":"5","key":"1032_CR8","doi-asserted-by":"publisher","first-page":"3637","DOI":"10.1152\/jn.00686.2005","volume":"94","author":"R Brette","year":"2005","unstructured":"Brette R, Gerstner W (2005) Adaptive exponential integrate-and-fire model as an effective description of neuronal activity. J Neurophysiol 94(5):3637\u20133642","journal-title":"J Neurophysiol"},{"issue":"3","key":"1032_CR9","doi-asserted-by":"publisher","first-page":"1208","DOI":"10.1152\/jn.1994.72.3.1208","volume":"72","author":"DE Brunn","year":"1994","unstructured":"Brunn DE, Dean J (1994) Intersegmental and local interneurons in the metathorax of the stick insect Carausius morosus that monitor middle leg position. J Neurophysiol 72(3):1208\u20131219","journal-title":"J Neurophysiol"},{"key":"1032_CR10","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/s00422-006-0068-6","volume":"95","author":"AN Burkitt","year":"2006","unstructured":"Burkitt AN (2006) A review of the integrate-and-fire neuron model: I. homogeneous synaptic input. Biol Cybern 95:1\u201319","journal-title":"Biol Cybern"},{"key":"1032_CR11","doi-asserted-by":"publisher","DOI":"10.1093\/acprof:oso\/9780198523444.001.0001","volume-title":"The neurobiology of an insect brain","author":"M Burrows","year":"1996","unstructured":"Burrows M (1996) The neurobiology of an insect brain. Oxford University Press"},{"issue":"1","key":"1032_CR12","doi-asserted-by":"publisher","first-page":"81","DOI":"10.1242\/jeb.144.1.81","volume":"144","author":"A B\u00fcschges","year":"1989","unstructured":"B\u00fcschges A (1989) Processing of sensory input from the femoral chordotonal organ by spiking interneurones of stick insects. J Exp Biol 144(1):81\u2013111","journal-title":"J Exp Biol"},{"issue":"2","key":"1032_CR13","doi-asserted-by":"publisher","first-page":"191","DOI":"10.1242\/jeb.42.2.191","volume":"42","author":"K Chapman","year":"1965","unstructured":"Chapman K (1965) Campaniform sensilla on the tactile spines of the legs of the cockroach. J Exp Biol 42(2):191\u2013203","journal-title":"J Exp Biol"},{"issue":"4868","key":"1032_CR14","doi-asserted-by":"publisher","first-page":"699","DOI":"10.1038\/197699a0","volume":"197","author":"K Chapman","year":"1963","unstructured":"Chapman K, Smith R (1963) A linear transfer function underlying impulse frequency modulation in a cockroach mechanoreceptor. Nature 197(4868):699\u2013700","journal-title":"Nature"},{"issue":"2","key":"1032_CR15","doi-asserted-by":"publisher","first-page":"149","DOI":"10.1007\/s004220050551","volume":"81","author":"JH Cocatre-Zilgien","year":"1999","unstructured":"Cocatre-Zilgien JH, Delcomyn F (1999) Modeling stress and strain in an insect leg for simulation of campaniform sensilla responses to external forces. Biol Cybern 81(2):149\u2013160","journal-title":"Biol Cybern"},{"issue":"9","key":"1032_CR16","doi-asserted-by":"publisher","first-page":"1617","DOI":"10.1242\/jeb.02166","volume":"209","author":"NJ Cowan","year":"2006","unstructured":"Cowan NJ, Lee J, Full RJ (2006) Task-level control of rapid wall following in the american cockroach. J Exp Biol 209(9):1617\u20131629","journal-title":"J Exp Biol"},{"key":"1032_CR17","doi-asserted-by":"publisher","first-page":"235","DOI":"10.1007\/BF00606541","volume":"112","author":"H Cruse","year":"1976","unstructured":"Cruse H (1976) The function of the legs in the free walking stick insect, Carausius morosus. J Comp Physiol 112:235\u2013262","journal-title":"J Comp Physiol"},{"key":"1032_CR18","doi-asserted-by":"publisher","first-page":"695","DOI":"10.1007\/BF01350223","volume":"154","author":"H Cruse","year":"1984","unstructured":"Cruse H, Dean J, Suilmann M (1984) The contributions of diverse sense organs to the control of leg movement by a walking insect. J Comp Physiol A 154:695\u2013705","journal-title":"J Comp Physiol A"},{"key":"1032_CR19","doi-asserted-by":"crossref","unstructured":"Cruse H, D\u00fcrr V, Schilling M, et al (2009) Principles of insect locomotion. In: Spatial temporal patterns for action-oriented perception in roving robots. Springer, p 43\u201396","DOI":"10.1007\/978-3-540-88464-4_2"},{"key":"1032_CR20","doi-asserted-by":"publisher","DOI":"10.1016\/j.cophys.2021.03.001","volume":"22","author":"CJ Dallmann","year":"2021","unstructured":"Dallmann CJ, Karashchuk P, Brunton BW et al (2021) A leg to stand on: computational models of proprioception. Curr Opin Physiol 22:100426","journal-title":"Curr Opin Physiol"},{"key":"1032_CR21","first-page":"49","volume-title":"Insect Locomotion Hamburg","author":"J Dean","year":"1985","unstructured":"Dean J (1985) A simulation of proprioceptive input from the coxal hair rows of the stick insect: possible effect of step velocity on the representation of joint angle. Insect Locomotion Hamburg. Paul Parey, pp 49\u201357"},{"key":"1032_CR22","first-page":"1","volume-title":"Advances in insect physiology","author":"LH Field","year":"1998","unstructured":"Field LH, Matheson T (1998) Chordotonal organs of insects. Advances in Insect Physiology 27. Elsevier, pp 1\u2013228"},{"key":"1032_CR23","first-page":"66","volume-title":"Breakthroughs in statistics: Methodology and distribution","author":"RA Fisher","year":"1970","unstructured":"Fisher RA (1970) Statistical methods for research workers. Breakthroughs in statistics: Methodology and distribution. Springer, pp 66\u201370"},{"issue":"6","key":"1032_CR24","doi-asserted-by":"publisher","first-page":"803","DOI":"10.1007\/BF00611597","volume":"155","author":"A French","year":"1984","unstructured":"French A (1984) Action potential adaptation in the femoral tactile spine of the cockroach, Periplaneta americana. J Comp Physiol A 155(6):803\u2013812","journal-title":"J Comp Physiol A"},{"key":"1032_CR25","doi-asserted-by":"publisher","first-page":"739","DOI":"10.1007\/s00359-002-0363-1","volume":"188","author":"AS French","year":"2002","unstructured":"French AS, Torkkeli PH, Seyfarth EA (2002) From stress and strain to spikes: mechanotransduction in spider slit sensilla. J Comp Physiol A 188:739\u2013752","journal-title":"J Comp Physiol A"},{"issue":"17","key":"1032_CR26","doi-asserted-by":"publisher","first-page":"3847","DOI":"10.1016\/j.cub.2022.07.005","volume":"32","author":"C Gebehart","year":"2022","unstructured":"Gebehart C, Hooper SL, B\u00fcschges A (2022) Non-linear multimodal integration in a distributed premotor network controls proprioceptive reflex gain in the insect leg. Curr Biol 32(17):3847\u20133854","journal-title":"Curr Biol"},{"issue":"13","key":"1032_CR27","doi-asserted-by":"publisher","first-page":"2265","DOI":"10.1242\/jeb.204.13.2265","volume":"204","author":"M Gebhardt","year":"2001","unstructured":"Gebhardt M, Honegger HW (2001) Physiological characterisation of antennal mechanosensory descending interneurons in an insect (gryllus bimaculatus, gryllus campestris) brain. J Exp Biol 204(13):2265\u20132275","journal-title":"J Exp Biol"},{"key":"1032_CR28","doi-asserted-by":"crossref","unstructured":"Gollin A, D\u00fcrr V (2018) Estimating body pitch from distributed proprioception in a hexapod. In: Biomimetic and Biohybrid Systems: 7th International Conference, Living Machines 2018, Paris, France, July 17\u201320, 2018, Proceedings 7, Springer, pp 187\u2013199","DOI":"10.1007\/978-3-319-95972-6_20"},{"issue":"2","key":"1032_CR29","doi-asserted-by":"publisher","first-page":"281","DOI":"10.1002\/jmor.1051030205","volume":"103","author":"A Hannah-Alava","year":"1958","unstructured":"Hannah-Alava A (1958) Morphology and chaetotaxy of the legs of drosophila melanogaster. J Morphol 103(2):281\u2013310","journal-title":"J Morphol"},{"issue":"1","key":"1032_CR30","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1111\/j.1365-3032.1986.tb00386.x","volume":"11","author":"T Hofmann","year":"1986","unstructured":"Hofmann T, B\u00e4ssler U (1986) Response characteristics of single trochanteral campaniform sensilla in the stick insect, cuniculina impigra. Physiol Entomol 11(1):17\u201321","journal-title":"Physiol Entomol"},{"issue":"1","key":"1032_CR31","doi-asserted-by":"publisher","first-page":"207","DOI":"10.1242\/jeb.114.1.207","volume":"114","author":"T Hofmann","year":"1985","unstructured":"Hofmann T, Koch U, B\u00e4ssler U (1985) Physiology of the femoral chordotonal organ in the stick insect, Cuniculina impigra. J Exp Biol 114(1):207\u2013223","journal-title":"J Exp Biol"},{"issue":"5","key":"1032_CR32","doi-asserted-by":"publisher","first-page":"1063","DOI":"10.1109\/TNN.2004.832719","volume":"15","author":"EM Izhikevich","year":"2004","unstructured":"Izhikevich EM (2004) Which model to use for cortical spiking neurons? IEEE Trans Neural Netw 15(5):1063\u20131070","journal-title":"IEEE Trans Neural Netw"},{"issue":"2","key":"1032_CR33","doi-asserted-by":"publisher","first-page":"398","DOI":"10.1152\/jn.00169.2021","volume":"126","author":"B Jaske","year":"2021","unstructured":"Jaske B, Lepreux G, D\u00fcrr V (2021) Input of hair field afferents to a descending interneuron. J Neurophysiol 126(2):398\u2013412","journal-title":"J Neurophysiol"},{"issue":"14","key":"1032_CR34","doi-asserted-by":"publisher","first-page":"4923","DOI":"10.1523\/JNEUROSCI.5777-11.2012","volume":"32","author":"PW Jones","year":"2012","unstructured":"Jones PW, Gabbiani F (2012) Logarithmic compression of sensory signals within the dendritic tree of a collision-sensitive neuron. J Neurosci 32(14):4923\u20134934","journal-title":"J Neurosci"},{"issue":"1","key":"1032_CR35","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1007\/BF00353956","volume":"44","author":"S Kemmerling","year":"1982","unstructured":"Kemmerling S, Varju D (1982) Regulation of the body-substrate-distance in the stick insect: step responses and modelling the control system. Biol Cybern 44(1):59\u201366","journal-title":"Biol Cybern"},{"issue":"1\u20132","key":"1032_CR36","doi-asserted-by":"publisher","first-page":"116","DOI":"10.1016\/j.jphysparis.2012.06.001","volume":"107","author":"AF Krause","year":"2013","unstructured":"Krause AF, Winkler A, D\u00fcrr V (2013) Central drive and proprioceptive control of antennal movements in the walking stick insect. J Physiol Paris 107(1\u20132):116\u2013129","journal-title":"J Physiol Paris"},{"key":"1032_CR37","first-page":"59","volume-title":"Advances in Insect Physiology","author":"A Krishnan","year":"2015","unstructured":"Krishnan A, Sane SP (2015) Antennal mechanosensors and their evolutionary antecedents. Advances in Insect Physiology 49. Elsevier, pp 59\u201399"},{"issue":"2","key":"1032_CR38","doi-asserted-by":"publisher","first-page":"510","DOI":"10.1111\/j.1749-6632.1961.tb35556.x","volume":"94","author":"WR Loewenstein","year":"1961","unstructured":"Loewenstein WR (1961) Excitation and inactivation in a receptor membrane. Ann N Y Acad Sci 94(2):510\u2013534","journal-title":"Ann N Y Acad Sci"},{"issue":"3","key":"1032_CR39","doi-asserted-by":"publisher","first-page":"636","DOI":"10.1016\/j.neuron.2018.09.009","volume":"100","author":"A Mamiya","year":"2018","unstructured":"Mamiya A, Gurung P, Tuthill JC (2018) Neural coding of leg proprioception in Drosophila. Neuron 100(3):636\u2013650","journal-title":"Neuron"},{"issue":"5","key":"1032_CR40","doi-asserted-by":"publisher","first-page":"475","DOI":"10.1007\/BF00342998","volume":"45","author":"H Markl","year":"1962","unstructured":"Markl H (1962) Borstenfelder an den Gelenken als Schweresinnesorgane bei Ameisen und anderen Hymenopteren. Z vgl Physiol 45(5):475\u2013569","journal-title":"Z vgl Physiol"},{"key":"1032_CR41","doi-asserted-by":"crossref","unstructured":"Mastella M, Chicca E (2021) A hardware-friendly neuromorphic spiking neural network for frequency detection and fine texture decoding. In: 2021 IEEE International Symposium on Circuits and Systems (ISCAS), IEEE, pp 1\u20135","DOI":"10.1109\/ISCAS51556.2021.9401377"},{"key":"1032_CR42","doi-asserted-by":"publisher","first-page":"509","DOI":"10.1007\/BF00191466","volume":"170","author":"T Matheson","year":"1992","unstructured":"Matheson T (1992) Range fractionation in the locust metathoracic femoral chordotonal organ. J Comp Physiol A 170:509\u2013520","journal-title":"J Comp Physiol A"},{"key":"1032_CR43","unstructured":"McIver SB (1985) Mechanoreception. In: Comprehensive insect physiology, biochemistry and pharmacology, vol\u00a02. Pergamon Oxford, chap\u00a02, p 71\u2013132"},{"key":"1032_CR44","unstructured":"Mill PJ (1976) Structure and Function of Proprioceptors in the Invertebrates. Chapman and Hall, London."},{"key":"1032_CR45","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1007\/s00422-008-0264-7","volume":"99","author":"R Naud","year":"2008","unstructured":"Naud R, Marcille N, Clopath C et al (2008) Firing patterns in the adaptive exponential integrate-and-fire model. Biol Cybern 99:335\u2013347","journal-title":"Biol Cybern"},{"issue":"11","key":"1032_CR46","doi-asserted-by":"publisher","first-page":"2397","DOI":"10.1242\/jeb.198.11.2397","volume":"198","author":"PL Newland","year":"1995","unstructured":"Newland PL, Watkins B, Emptage NJ, Nagayama T\u00a0(1995) The structure, response properties and development of a hair plate on the mesothoracic leg of the locust. J Exp Biol 198(11):2397\u20132404","journal-title":"J Exp Biol"},{"issue":"24","key":"1032_CR47","doi-asserted-by":"publisher","first-page":"4301","DOI":"10.1242\/jeb.204.24.4301","volume":"204","author":"J Okada","year":"2001","unstructured":"Okada J, Toh Y (2001) Peripheral representation of antennal orientation by the scapal hair plate of the cockroach Periplaneta americana. J Exp Biol 204(24):4301\u20134309","journal-title":"J Exp Biol"},{"issue":"4","key":"1032_CR48","doi-asserted-by":"publisher","first-page":"467","DOI":"10.1242\/jeb.15.4.467","volume":"15","author":"J Pringle","year":"1938","unstructured":"Pringle J (1938) Proprioception in insects: III. the function of the hair sensilla at the joints. J Exp Biol 15(4):467\u2013473","journal-title":"J Exp Biol"},{"key":"1032_CR49","unstructured":"Schmitz J (1985a) Control of the leg joints in stick insects: differences in the reflex properties between the standing and the walking states. Insect locomotion pp 27\u201332"},{"key":"1032_CR50","unstructured":"Schmitz J (1985b) Systemanalytische Untersuchungen zur Positionsregelung proximaler Beingelenke der Stabheuschrecke (Carausius morosus): Eigenschaften des Coxa-Trochanter Regelkreises im stehenden und laufenden Tier. Doctoral Dissertation, Bielefeld University."},{"issue":"1","key":"1032_CR51","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1007\/BF00363976","volume":"55","author":"J Schmitz","year":"1986","unstructured":"Schmitz J (1986) Properties of the feedback system controlling the coxa-trochanter joint in the stick insect Carausius morosus. Biol Cybern 55(1):35\u201342","journal-title":"Biol Cybern"},{"issue":"4","key":"1032_CR52","doi-asserted-by":"publisher","first-page":"424","DOI":"10.1002\/(SICI)1097-4695(200003)42:4<424::AID-NEU4>3.0.CO;2-0","volume":"42","author":"J Schmitz","year":"2000","unstructured":"Schmitz J, Stein W (2000) Convergence of load and movement information onto leg motoneurons in insects. J Neurobiol 42(4):424\u2013436","journal-title":"J Neurobiol"},{"issue":"1","key":"1032_CR53","doi-asserted-by":"publisher","first-page":"512","DOI":"10.1152\/jn.1999.82.1.512","volume":"82","author":"W Stein","year":"1999","unstructured":"Stein W, Schmitz J (1999) Multimodal convergence of presynaptic afferent inhibition in insect proprioceptors. J Neurophysiol 82(1):512\u2013514","journal-title":"J Neurophysiol"},{"issue":"6","key":"1032_CR54","doi-asserted-by":"publisher","DOI":"10.1088\/1748-3190\/ac1ced","volume":"16","author":"NS Szczecinski","year":"2021","unstructured":"Szczecinski NS, Dallmann CJ, Quinn RD et al (2021) A computational model of insect campaniform sensilla predicts encoding of forces during walking. Bioinspir Biomim 16(6):065001","journal-title":"Bioinspir Biomim"},{"issue":"12","key":"1032_CR55","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0085321","volume":"8","author":"LM Theunissen","year":"2013","unstructured":"Theunissen LM, D\u00fcrr V (2013) Insects use two distinct classes of steps during unrestrained locomotion. PLoS One 8(12):e85321","journal-title":"PLoS One"},{"issue":"18","key":"1032_CR56","first-page":"3242","volume":"217","author":"LM Theunissen","year":"2014","unstructured":"Theunissen LM, Vikram S, D\u00fcrr V (2014) Spatial co-ordination of foot contacts in unrestrained climbing insects. J Exp Biol 217(18):3242\u20133253","journal-title":"J Exp Biol"},{"issue":"3","key":"1032_CR57","first-page":"340","volume":"218","author":"LM Theunissen","year":"2015","unstructured":"Theunissen LM, Bekemeier HH, D\u00fcrr V (2015) Comparative whole-body kinematics of closely related insect species with different body morphology. J Exp Biol 218(3):340\u2013352","journal-title":"J Exp Biol"},{"key":"1032_CR58","doi-asserted-by":"crossref","unstructured":"Thurm U (1965) An insect mechanoreceptor. Part I: Fine structure and adequate stimulus. In: Cold Spring Harbor symposia on quantitative biology, Cold Spring Harbor Laboratory Press, pp 75\u201382","DOI":"10.1101\/SQB.1965.030.01.011"},{"key":"1032_CR59","doi-asserted-by":"publisher","DOI":"10.2307\/3001913","author":"JW Tukey","year":"1949","unstructured":"Tukey JW (1949) Comparing individual means in the analysis of variance. Biometrics. https:\/\/doi.org\/10.2307\/3001913","journal-title":"Biometrics"},{"issue":"5","key":"1032_CR60","doi-asserted-by":"publisher","first-page":"R194","DOI":"10.1016\/j.cub.2018.01.064","volume":"28","author":"JC Tuthill","year":"2018","unstructured":"Tuthill JC, Azim E (2018) Proprioception. Curr Biol 28(5):R194\u2013R203","journal-title":"Curr Biol"},{"issue":"20","key":"1032_CR61","doi-asserted-by":"publisher","first-page":"R1022","DOI":"10.1016\/j.cub.2016.06.070","volume":"26","author":"JC Tuthill","year":"2016","unstructured":"Tuthill JC, Wilson RI (2016) Mechanosensation and adaptive motor control in insects. Curr Biol 26(20):R1022\u2013R1038","journal-title":"Curr Biol"},{"key":"1032_CR62","doi-asserted-by":"crossref","unstructured":"van der Veen T, D\u00fcrr V, Chicca E (2026) Encoding of movement primitives and body posture through distributed proprioception in walking and climbing insects. Biological Cybernetics 120:3 https:\/\/doi.org\/10.1007\/s00422-025-01029-x","DOI":"10.1007\/s00422-025-01029-x"},{"key":"1032_CR63","doi-asserted-by":"publisher","DOI":"10.1016\/j.cobeha.2024.101459","volume":"60","author":"S Virdi","year":"2024","unstructured":"Virdi S, Sane SP (2024) Form, function and mechanics of femoral chordotonal organs in insects. Curr Opin Behav Sci 60:101459","journal-title":"Curr Opin Behav Sci"},{"key":"1032_CR64","doi-asserted-by":"publisher","first-page":"198","DOI":"10.1007\/BF00297860","volume":"48","author":"G Wendler","year":"1964","unstructured":"Wendler G (1964) Laufen und Stehen der Stabheuschrecke Carausius morosus: Sinnesborstenfelder in den Beingelenken als Glieder von Regelkreisen. Z vergl Physiol 48:198\u2013250","journal-title":"Z vergl Physiol"},{"issue":"1","key":"1032_CR65","doi-asserted-by":"publisher","first-page":"233","DOI":"10.1242\/jeb.64.1.233","volume":"64","author":"R Wong","year":"1976","unstructured":"Wong R, Pearson K (1976) Properties of the trochanteral hair plate and its function in the control of walking in the cockroach. J Exp Biol 64(1):233\u2013249","journal-title":"J Exp Biol"},{"issue":"7","key":"1032_CR66","doi-asserted-by":"publisher","DOI":"10.3390\/brainsci12070863","volume":"12","author":"K Yamazaki","year":"2022","unstructured":"Yamazaki K, Vo-Ho VK, Bulsara D et al (2022) Spiking neural networks and their applications: a review. Brain Sci 12(7):863","journal-title":"Brain Sci"},{"key":"1032_CR67","doi-asserted-by":"crossref","unstructured":"Zadokha B, Szczecinski NS (2024) Encoding 3d leg kinematics using spatially-distributed, population coded network model. In: Conference on Biomimetic and Biohybrid Systems, Springer, pp 317\u2013332","DOI":"10.1007\/978-3-031-72597-5_22"},{"issue":"3","key":"1032_CR68","doi-asserted-by":"publisher","first-page":"273","DOI":"10.1016\/j.asd.2004.05.005","volume":"33","author":"S Zill","year":"2004","unstructured":"Zill S, Schmitz J, B\u00fcschges A (2004) Load sensing and control of posture and locomotion. Arthropod Struct Dev 33(3):273\u2013286","journal-title":"Arthropod Struct Dev"},{"issue":"1","key":"1032_CR69","doi-asserted-by":"publisher","first-page":"435","DOI":"10.1242\/jeb.116.1.435","volume":"116","author":"SN Zill","year":"1985","unstructured":"Zill SN (1985) Plasticity and proprioception in insects. I. Responses and cellular properties of individual receptors of the locust metathoracic femoral chordotonal organ. J Exp Biol 116(1):435\u2013461","journal-title":"J Exp Biol"}],"container-title":["Biological Cybernetics"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00422-025-01032-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s00422-025-01032-2","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00422-025-01032-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,5,8]],"date-time":"2026-05-08T07:03:15Z","timestamp":1778223795000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s00422-025-01032-2"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,2,28]]},"references-count":69,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2026,4]]}},"alternative-id":["1032"],"URL":"https:\/\/doi.org\/10.1007\/s00422-025-01032-2","relation":{},"ISSN":["1432-0770"],"issn-type":[{"value":"1432-0770","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,2,28]]},"assertion":[{"value":"26 May 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"30 December 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"28 February 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare no competing interests.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"6"}}