Predictions not commands: active inference in the motor system

Rick A. Adams1, Stewart Shipp2, Karl J. Friston1
1The Wellcome Trust Centre for Neuroimaging, Institute of Neurology, University College London, London, UK
2UCL Institute of Ophthalmology, University College London, London, UK

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Akatsuka K, Wasaka T, Nakata H, Kida T, Kakigi R (2007) The effect of stimulus probability on the somatosensory mismatch field. Exp Brain Res 181:607–614

Andersen P, Raastad M, Storm JF (1990) Excitatory synaptic integration in hippocampal pyramids and dentate granule cells. Cold Spring Harb Symp Quant Biol 55:81–86

Angelucci A, Bullier J (2003) Reaching beyond the classical receptive field of V1 neurons: horizontal or feedback axons? J Physiol Paris 97:141–154

Arikuni T, Watanabe K, Kubota K (1988) Connections of area 8 with area 6 in the brain of the macaque monkey. J Comp Neurol 277:21–40

Asanuma H, Zarzecki P, Jankowska E, Hongo T, Marcus S (1979) Projection of individual pyramidal tract neurons to lumbar motor nuclei of the monkey. Exp Brain Res 34:73–89

Asanuma H, Larsen K, Yumiya H (1980) Peripheral input pathways to the monkey motor cortex. Exp Brain Res 38:349–355

Asanuma C, Thach WT, Jones EG (1983) Distribution of cerebellar terminations and their relation to other afferent terminations in the ventral lateral thalamic region of the monkey. Brain Res 286:237–265

Avermann M, Tomm C, Mateo C, Gerstner W, Petersen CCH (2012) Microcircuits of excitatory and inhibitory neurons in layer 2/3 of mouse barrel cortex. J Neurophysiol 107:3116–3134

Bannatyne BA, Edgley SA, Hammar I, Jankowska E, Maxwell DJ (2006) Differential projections of excitatory and inhibitory dorsal horn interneurons relaying information from group II muscle afferents in the cat spinal cord. J Neurosci 26:2871–2880

Barbas H (1986) Pattern in the laminar origin of corticocortical connections. J Comp Neurol. 252:415–422

Barbas H, Pandya DN (1987) Architecture and frontal cortical connections of the premotor cortex (area 6) in the rhesus monkey. J Comp Neurol 256:211–228

Barone P, Batardiere A, Knoblauch K, Kennedy H (2000) Laminar distribution of neurons in extrastriate areas projecting to visual areas V1 and V4 correlates with the hierarchical rank and indicates the operation of a distance rule. J Neurosci 20:3263–3281

Beck O, Chistiakova M, Obermayer K, Volgushev M (2005) Adaptation at synaptic connections to layer 2/3 pyramidal cells in rat visual cortex. J Neurophysiol 94:363–376

Bentivoglio M, Rustioni A (1986) Corticospinal neurons with branching axons to the dorsal column nuclei in the monkey. J Comp Neurol 253:260–276

Bernier P-M, Chua R, Bard C, Franks IM (2006) Updating of an internal model without proprioception: a deafferentation study. NeuroReport 17:1421–1425

Binshtok AM, Fleidervish IA, Sprengel R, Gutnick MJ (2006) NMDA receptors in layer 4 spiny stellate cells of the mouse barrel cortex contain the NR2C subunit. J Neurosci 26:708–715

Blakemore S-J, Sirigu A (2003) Action prediction in the cerebellum and in the parietal lobe. Exp Brain Res 153:239–245

Blakemore SJ, Wolpert D, Frith C (2000) Why can’t you tickle yourself? NeuroReport 11:R11–R16

Borra E, Belmalih A, Calzavara R, Gerbella M, Murata A, Rozzi S, Luppino G (2008) Cortical connections of the macaque anterior intraparietal (AIP) area. Cereb Cortex 18:1094–1111

Borst JG (2010) The low synaptic release probability in vivo. Trends Neurosci 33:259–266

Boudrias M-H, McPherson RL, Frost SB, Cheney PD (2010) Output properties and organization of the forelimb representation of motor areas on the lateral aspect of the hemisphere in rhesus macaques. Cereb Cortex 20:169–186

Branco T, Staras K (2009) The probability of neurotransmitter release: variability and feedback control at single synapses. Nat Rev Neurosci 10:373–383

Brodmann K (1909/1994) Localisation in the cerebral cortex [Vergleichende Lokalisationslehre der Großhirnrinde, 1909 (Trans: Garey LJ)]. Smith-Gordon, London

Brown P (1994) Pathophysiology of spasticity. J Neurol Neurosurg Psychiatr 57:773–777

Brown RG, Marsden CD (1988) Internal versus external cues and the control of attention in Parkinson’s disease. Brain 111:323–345

Brown H, Friston K, Bestmann S (2011) Active inference, attention, and motor preparation. Front Psychol 2:218

Büchel C, Friston KJ (1997) Modulation of connectivity in visual pathways by attention: cortical interactions evaluated with structural equation modelling and fMRI. Cereb Cortex 7:768–778

Bullier J, Hupé JM, James AC, Girard P (2001) The role of feedback connections in shaping the responses of visual cortical neurons. Prog Brain Res 134:193–204

Bullier J, Kennedy H, Salinger W (1984) Branching and laminar origin of projections between visual cortical areas in the cat. J Comp Neurol 228:329–341

Burton H, Fabri M (1995) Ipsilateral intracortical connections of physiologically defined cutaneous representations in areas 3b and 1 of macaque monkeys: projections in the vicinity of the central sulcus. J Comp Neurol 355:508–538

Burton H, Sinclair R (1996) Somatosensory cortex and tactile perceptions. In: Kruger L (ed) Touch and pain. Academic Press, London, pp 105–177

Buys EJ, Lemon RN, Mantel GW, Muir RB (1986) Selective facilitation of different hand muscles by single corticospinal neurones in the conscious monkey. J Physiol (Lond) 381:529–549

Campbell AW (1905) Histological studies on the localisation of cerebral function. Cambridge University Press, Cambridge, UK

Censor N, Sagi D, Cohen LG (2012) Common mechanisms of human perceptual and motor learning. Nat Rev Neurosci 13:658–664

Cheema S, Rustioni A, Whitsel BL (1985) Sensorimotor cortical projections to the primate cuneate nucleus. J Comp Neurol 240:196–211

Chen CC, Henson RN, Stephan KE, Kilner JM, Friston KJ (2009) Forward and backward connections in the brain: a DCM study of functional asymmetries. Neuroimage 45:453–462

Cheney PD, Fetz EE (1984) Corticomotoneuronal cells contribute to long-latency stretch reflexes in the rhesus monkey. J Physiol (Lond) 349:249–272

Cherian A, Krucoff MO, Miller LE (2011) Motor cortical prediction of EMG: evidence that a kinetic brain–machine interface may be robust across altered movement dynamics. J Neurophysiol 106:564–575

Clarke RJ, Johnson JW (2006) NMDA receptor NR2 subunit dependence of the slow component of magnesium unblock. J Neurosci 26:5825–5834

Covic EN, Sherman SM (2011) Synaptic properties of connections between the primary and secondary auditory cortices in mice. Cereb Cortex 21:2425–2441

Craig ADB (2008) Retrograde analyses of spinothalamic projections in the macaque monkey: input to the ventral lateral nucleus. J Comp Neurol 508:315–328

Cull-Candy SG, Leszkiewicz DN (2004) Role of distinct NMDA receptor subtypes at central synapses. Sci STKE 2004:re16

Cullen KE (2004) Sensory signals during active versus passive movement. Curr Opin Neurobiol 14:698–706

Dancause N, Barbay S, Frost SB, Plautz EJ, Stowe AM, Friel KM, Nudo RJ (2006) Ipsilateral connections of the ventral premotor cortex in a new world primate. J Comp Neurol 495:374–390

Darian-Smith C, Darian-Smith I, Burman K, Ratcliffe N (1993) Ipsilateral cortical projections to areas 3a, 3b, and 4 in the macaque monkey. J Comp Neurol 335:200–213

de Noordhout AM, Rapisarda G, Bogacz D, Gerard P, De Pasqua V, Pennisi G, Delwaide PJ (1999) Corticomotoneuronal synaptic connections in normal man: an electrophysiological study. Brain 122:1327–1340

De Pasquale R, Sherman SM (2011) Synaptic properties of corticocortical connections between the primary and secondary visual cortical areas in the mouse. J Neurosci 31:16494–16506

Deacon TW (1992) Cortical connections of the inferior arcuate sulcus cortex in the macaque brain. Brain Res 573:8–26

den Ouden HEM, Daunizeau J, Roiser J, Friston K, Stephan KE (2010) Striatal prediction error modulates cortical coupling. J Neurosci 30:3210–3219

Dietz V, Sinkjaer T (2007) Spastic movement disorder: impaired reflex function and altered muscle mechanics. Lancet Neurol 6:725–733

Dobrunz LE, Stevens CF (1997) Heterogeneity of release probability, facilitation, and depletion at central synapses. Neuron 18:995–1008

Dum RP, Strick PL (2005) Frontal lobe inputs to the digit representations of the motor areas on the lateral surface of the hemisphere. J Neurosci 25:1375–1386

Durstewitz D (2009) Implications of synaptic biophysics for recurrent network dynamics and active memory. Neural Netw 22:1189–1200

Eickhoff SB, Rottschy C, Zilles K (2007) Laminar distribution and co-distribution of neurotransmitter receptors in early human visual cortex. Brain Struct Funct 212:255–267

Feldman AG (2008) Threshold position control signifies a common spatial frame of reference for motor action and kinesthesia. Brain Res Bull 75:497–499

Feldman AG (2009) New insights into action–perception coupling. Exp Brain Res 194:39–58

Feldman H, Friston KJ (2010) Attention, uncertainty, and free-energy. Front Hum Neurosci 4:215

Feldman AG, Levin MF (2009) The equilibrium point hypothesis: past, present and future. Adv Exp Med Biol 629:699–726

Feldman AG, Orlovsky GN (1972) The influence of different descending systems on the tonic stretch reflex in the cat. Exp Neurol 37:481–494

Felleman DJ, Van Essen DC (1991) Distributed hierarchical processing in the primate cerebral cortex. Cereb Cortex 1:1–47

Ferrer JM, Kato N, Price DJ (1992) Organization of association projections from area 17 to areas 18 and 19 and to suprasylvian areas in the cat’s visual cortex. J Comp Neurol 316:261–278

Fox K, Sato H, Daw N (1990) The effect of varying stimulus intensity on NMDA-receptor activity in cat visual cortex. J Neurophysiol 64:1413–1428

Friston KJ (2003) Learning and inference in the brain. Neural Netw 16:1325–1352

Friston KJ (2005) A theory of cortical responses. Philos Trans R Soc Lond B Biol Sci 360:815–836

Friston KJ (2010) The free-energy principle: a unified brain theory? Nat Rev Neurosci 11:127–138

Friston KJ (2011) What is optimal about motor control? Neuron 72:488–498

Friston KJ, Kiebel S (2009) Cortical circuits for perceptual inference. Neural Netw 22:1093–1104

Friston KJ, Kilner J, Harrison L (2006) A free energy principle for the brain. J Physiol Paris 100:70–87

Friston KJ, Daunizeau J, Kilner J, Kiebel SJ (2010) Action and behavior: a free-energy formulation. Biol Cybern 102:227–260

Friston KJ, Mattout J, Kilner J (2011) Action understanding and active inference. Biol Cybern 104:137–160

Friston KJ, Shiner T, Fitzgerald T, Galea JM, Adams R, Brown H, Dolan RJ, Moran R, Stephan KE, Bestmann S (2012) Dopamine, affordance and active inference. PLoS Comput Biol 8:e1002327

Galea MP, Darian-Smith I (1994) Multiple corticospinal neuron populations in the macaque monkey are specified by their unique cortical origins, spinal terminations, and connections. Cereb Cortex 4:166–194

Garrido MI, Kilner JM, Stephan KE, Friston KJ (2009) The mismatch negativity: a review of underlying mechanisms. Clin Neurophysiol 120:453–463

Gentilucci M, Toni I, Chieffi S, Pavesi G (1994) The role of proprioception in the control of prehension movements: a kinematic study in a peripherally deafferented patient and in normal subjects. Exp Brain Res 99:483–500

Gerbella M, Belmalih A, Borra E, Rozzi S, Luppino G (2011) Cortical connections of the anterior (F5a) subdivision of the macaque ventral premotor area F5. Brain Struct Func 216:43–65

Geyer S, Matelli M, Luppino G, Zilles K (2000) Functional neuroanatomy of the primate isocortical motor system. Anat Embryol 202:443–474

Ghosh S, Porter R (1988) Corticocortical synaptic influences on morphologically identified pyramidal neurones in the motor cortex of the monkey. J Physiol (Lond) 400:617–629

Ghosh S, Brinkman C, Porter R (1987) A quantitative study of the distribution of neurons projecting to the precentral motor cortex in the monkey (M. fascicularis). J Comp Neurol 259:424–444

Ginanneschi F, Del Santo F, Dominici F, Gelli F, Mazzocchio R, Rossi A (2005) Changes in corticomotor excitability of hand muscles in relation to static shoulder positions. Exp Brain Res 161:374–382

Godschalk M, Lemon RN, Kuypers HG, Ronday HK (1984) Cortical afferents and efferents of monkey postarcuate area: an anatomical and electrophysiological study. Exp Brain Res 56:410–424

Granit R (1955) Receptors and sensory perception. Yale University Press, Newhaven

Graziano M (2006) The organization of behavioral repertoire in motor cortex. Annu Rev Neurosci 29:105–134

Griffin DM, Hudson HM, Belhaj-Saif A, Cheney PD (2011) Hijacking cortical motor output with repetitive microstimulation. J Neurosci 31:13088–13096

Groh A, de Kock CP, Wimmer VC, Sakmann B, Kuner T (2008) Driver or coincidence detector: modal switch of a corticothalamic giant synapse controlled by spontaneous activity and short-term depression. J Neurosci 28:9652–9663

Gurney K, Prescott TJ, Redgrave P (2001) A computational model of action selection in the basal ganglia. I A new functional anatomy. Biol Cybern 84:401–410

Hantman AW, Jessell TM (2010) Clarke’s column neurons as the focus of a corticospinal corollary circuit. Nat Neurosci 13:1233–1239

Harrison SA, Tong F (2009) Decoding reveals the contents of visual working memory in early visual areas. Nature 458:632–635

Harrison LM, Stephan KE, Rees G, Friston KJ (2007) Extra-classical receptive field effects measured in striate cortex with fMRI. Neuroimage 34:1199–1208

Hatanaka N, Nambu A, Yamashita A, Takada M, Tokuno H (2001) Somatotopic arrangement and corticocortical inputs of the hindlimb region of the primary motor cortex in the macaque monkey. Neurosci Res 40:9–22

Hatanaka N, Tokuno H, Hamada I, Inase M, Ito Y, Imanishi M, Hasegawa N, Akazawa T, Nambu A, Takada M (2003) Thalamocortical and intracortical connections of monkey cingulate motor areas. J Comp Neurol 462:121–138

Hatsopoulos NG, Suminski AJ (2011) Sensing with the motor cortex. Neuron 72:477–487

Heckman CJ, Hyngstrom AS, Johnson MD (2008) Active properties of motoneurone dendrites: diffuse descending neuromodulation, focused local inhibition. J Physiol (Lond) 586:1225–1231

Helmholtz H (1860/1962) Handbuch der physiologischen Optik, vol 3. (English Trans: Southall JPC, ed). New York, Dover

Helmich RC, Hallett M, Deuschl G, Toni I, Bloem BR (2012) Cerebral causes and consequences of Parkinsonian resting tremor: a tale of two circuits? Brain. doi: 10.1093/brain/aws023

Henson RN, Gagnepain P (2010) Predictive, interactive multiple memory systems. Hippocampus 20:1315–1326

Herter TM, Korbel T, Scott SH (2009) Comparison of neural responses in primary motor cortex to transient and continuous loads during posture. J Neurophysiol 101:150–163

Hilgetag CC, Grant S (2010) Cytoarchitectural differences are a key determinant of laminar projection origins in the visual cortex. Neuroimage 51:1006–1017

Hirai T, Jones EG (1988) Segregation of lemniscal inputs and motor cortex outputs in cat ventral thalamic nuclei: application of a novel technique. Exp Brain Res 71:329–344

Horne MK, Tracey DJ (1979) The afferents and projections of the ventroposterolateral thalamus in the monkey. Exp Brain Res 36:129–141

Hsu CL, Yang HW, Yen CT, Min MY (2010) Comparison of synaptic transmission and plasticity between sensory and cortical synapses on relay neurons in the ventrobasal nucleus of the rat thalamus. J Physiol (Lond) 588:4347–4363

Huerta MF, Pons TP (1990) Primary motor cortex receives input from area 3a in macaques. Brain Res 537:367–371

Huffman KJ, Krubitzer L (2001) Thalamo-cortical connections of areas 3a and M1 in marmoset monkeys. J Comp Neurol 435:291–310

Hultborn H, Meunier S, Pierrot-Deseilligny E, Shindo M (1987) Changes in presynaptic inhibition of Ia fibres at the onset of voluntary contraction in man. J Physiol 389:757–772

Hummelsheim H, Bianchetti M, Wiesendanger M, Wiesendanger R (1988) Sensory inputs to the agranular motor fields: a comparison between precentral, supplementary-motor and premotor areas in the monkey. Exp Brain Res 69:289–298

Hupé JM, James AC, Payne BR, Lomber SG, Girard P, Bullier J (1998) Cortical feedback improves discrimination between figure and background by V1, V2 and V3 neurons. Nature 394:784–787

Jankowska E, Padel Y, Tanaka R (1975) The mode of activation of pyramidal tract cells by intracortical stimuli. J Physiol 249:617–636

Johnson PB, Ferraina S (1996) Cortical networks for visual reaching: intrinsic frontal lobe connectivity. Eur J Neurosci 8:1358–1362

Jones EG, Coulter JD, Hendry SH (1978) Intracortical connectivity of architectonic fields in the somatic sensory, motor and parietal cortex of monkeys. J Comp Neurol 181:291–347

Kalil K (1981) Projections of the cerebellar and dorsal column nuclei upon the thalamus of the rhesus monkey. J Comp Neurol 195:25–50

Kasser RJ, Cheney PD (1985) Characteristics of corticomotoneuronal postspike facilitation and reciprocal suppression of EMG activity in the monkey. J Neurophysiol 53:959–978

Kiebel SJ, Klöppel S, Weiskopf N, Friston KJ (2007) Dynamic causal modeling: a generative model of slice timing in fMRI. Neuroimage 34:1487–1496

Krakauer JW, Ghilardi MF, Ghez C (1999) Independent learning of internal models for kinematic and dynamic control of reaching. Nat Neurosci 2:1026–1031

Künzle H (1978a) An autoradiographic analysis of the efferent connections from premotor and adjacent prefrontal regions (areas 6 and 9) in Macaca fascicularis. Brain Behav Evol 15:185–234

Künzle H (1978b) Cortico-cortical efferents of primary motor and somatosensory regions of the cerebral cortex in Macaca fascicularis. Neuroscience 3:25–39

Kurata K (1991) Corticocortical inputs to the dorsal and ventral aspects of the premotor cortex of macaque monkeys. Neurosci Res 12:263–280

Kurtzer IL, Pruszynski JA, Scott SH (2008) Long-latency reflexes of the human arm reflect an internal model of limb dynamics. Curr Biol 18:449–453

Kuypers HGJM (1981) Anatomy of the descending pathways. In: Brookhart J, Mountcastle V (eds) Handbook of physiology—the nervous system II. American Physiological Society, Bethesda, MD, pp 597–666

Lamotte d’Incamps B, Ascher P (2008) Four excitatory postsynaptic ionotropic receptors coactivated at the motoneuron–Renshaw cell synapse. J Neurosci 28:14121–14131

Larkum ME, Zhu JJ, Sakmann B (1999) A new cellular mechanism for coupling inputs arriving at different cortical layers. Nature 398:338–341

Larkum ME, Senn W, Lüscher H-R (2004) Top-down dendritic input increases the gain of layer 5 pyramidal neurons. Cereb Cortex 14:1059–1070

Larkum ME, Nevian T, Sandler M, Polsky A, Schiller J (2009) Synaptic integration in tuft dendrites of layer 5 pyramidal neurons: a new unifying principle. Science 325:756–760

Lee CC, Sherman SM (2008) Synaptic properties of thalamic and intracortical inputs to layer 4 of the first- and higher-order cortical areas in the auditory and somatosensory systems. J Neurophysiol 100:317–326

Leichnetz GR (1986) Afferent and efferent connections of the dorsolateral precentral gyrus (area 4, hand/arm region) in the macaque monkey, with comparisons to area 8. J Comp Neurol 254:460–492

Leichnetz GR (2001) Connections of the medial posterior parietal cortex (area 7m) in the monkey. Anat Rec 263:215–236

Lemon RN (2008) Descending pathways in motor control. Ann Rev Neurosci 31:195–218

Lemon RN, Griffiths J (2005) Comparing the function of the corticospinal system in different species: organizational differences for motor specialization? Muscle Nerve 32:261–279

Lemon RN, Porter R (1976) Afferent input to movement-related precentral neurones in conscious monkeys. Proc R Soc Lond B Biol Sci 194:313–339

Lemon RN, van der Burg J (1979) Short-latency peripheral inputs to thalamic neurones projecting to the motor cortex in the monkey. Exp Brain Res 36:445–462

Li J, Guido W, Bickford ME (2003) Two distinct types of corticothalamic EPSPs and their contribution to short-term synaptic plasticity. J Neurophysiol 90:3429–3440

Lindström S (1973) Recurrent control from motor axon collaterals of Ia inhibitory pathways in the spinal cord of the cat. Acta Physiol Scand Suppl 392:1–43

Lu MT, Preston JB, Strick PL (1994) Interconnections between the prefrontal cortex and the premotor areas in the frontal lobe. J Comp Neurol 341:375–392

Luppino G, Matelli M, Camarda R, Rizzolatti G (1993) Corticocortical connections of area F3 (SMA-proper) and area F6 (pre-SMA) in the macaque monkey. J Comp Neurol 338:114–140

Macmillan M (2012) Alfred Walter Campbell and the visual functions of the occipital cortex. Cortex (in press)

Macchi G, Jones EG (1997) Toward an agreement on terminology on nuclear and subnuclear divisions of the motor thalamus. J Neurosurg 86:670–685

Martinez-Conde S, Cudeiro J, Grieve KL, Rodriguez R, Rivadulla C, Acuna C (1999) Effects of feedback projections from area 18 layers 2/3 to area 17 layers 2/3 in the cat visual cortex. J Neurophysiol 82:2667–2675

Matelli M, Camarda R, Glickstein M, Rizzolatti G (1986) Afferent and efferent projections of the inferior area 6 in the macaque monkey. J Comp Neurol 251:281–298

Matthews PB (1959) A study of certain factors influencing the stretch reflex of the decerebrate cat. J Physiol 147:547–564

Matthews PB (1991) The human stretch reflex and the motor cortex. Trends Neurosci 14:87–91

Matyas F, Sreenivasan V, Marbach F, Wacongne C, Barsy B, Mateo C, Aronoff R, Petersen CCH (2010) Motor control by sensory cortex. Science 330:1240–1243

McCulloch RM, Johnston GAR, Game CJA, Curtis DR (1974) The differential sensitivity of spinal interneurones and Renshaw cells to kainate and N-methyl-d-aspartate. Exp Brain Res 21:515–518

McKiernan BJ, Marcario JK, Karrer JH, Cheney PD (1998) Corticomotoneuronal postspike effects in shoulder, elbow, wrist, digit, and intrinsic hand muscles during a reach and prehension task. J Neurophysiol 80:1961–1980

McNally GP, Johansen JP, Blair HT (2011) Placing prediction into the fear circuit. Trends Neurosci 34:283–292

Medalla M, Barbas H (2009) Synapses with inhibitory neurons differentiate anterior cingulate from dorsolateral prefrontal pathways associated with cognitive control. Neuron 61:609–620

Merton PA (1953) Speculations on the servo-control of movement. In: Wolstenholme GEW (ed) The spinal cord. Churchill, London, pp 247–255

Mottram CJ, Suresh NL, Heckman CJ, Gorassini MA, Rymer WZ (2009) Origins of abnormal excitability in biceps brachii motoneurons of spastic-paretic stroke survivors. J Neurophysiol 102:2026–2038

Muakkassa KF, Strick PL (1979) Frontal lobe inputs to primate motor cortex: evidence for four somatotopically organized “premotor” areas. Brain Res 177:176–182

Muckli L, Kohler A, Kriegeskorte N, Singer W (2005) Primary visual cortex activity along the apparent-motion trace reflects illusory perception. PLoS Biol 3:e265

Mulliken GH, Musallam S, Andersen RA (2008) Forward estimation of movement state in posterior parietal cortex. Proc Natl Acad Sci USA 105:8170–8177

Mumford D (1992) On the computational architecture of the neocortex. II. The role of cortico-cortical loops. Biol Cybern 66:241–251

Mumford D (1994) Neuronal architectures for pattern-theoretic problems. In: Koch C, Davis JL (eds) Large-scale neuronal theories of the brain. MIT Press, Cambridge, MA, pp 125–152

Muñoz A, Woods TM, Jones EG (1999) Laminar and cellular distribution of AMPA, kainate, and NMDA receptor subunits in monkey sensory-motor cortex. J Comp Neurol 407:472–490

Murray SO, Kersten D, Olshausen BA, Schrater P, Woods DL (2002) Shape perception reduces activity in human primary visual cortex. Proc Natl Acad Sci USA 99:15164–15169

Murray SO, Olman CA, Kersten D (2006) Spatially specific FMRI repetition effects in human visual cortex. J Neurophysiol 95:2439–2445

Mutel V, Buchy D, Kilngelschmidt A, Messer J, Bleuel Z, Kemp JA, Grayson Richards J (1998) In vitro binding properties in rat brain of [3H]Ro 25–6981, a potent and selective antagonist of NMDA receptors containing NR2B subunits. J Neurochem 70:2147–2155

Olsen SR, Bortone DS, Adesnik H, Scanziani M (2012) Gain control by layer six in cortical circuits of vision. Nature 483:47–52

Padberg J, Cerkevich C, Engle J, Rajan AT, Recanzone G, Kaas J, Krubitzer L (2009) Thalamocortical connections of parietal somatosensory cortical fields in macaque monkeys are highly divergent and convergent. Cereb Cortex 19:2038–2064

Palecek JI, Abdrachmanova G, Vlachová V, Vyklick L Jr (1999) Properties of NMDA receptors in rat spinal cord motoneurons. Eur J Neurosci 11:827–836

Palmer E, Ashby P (1992) Corticospinal projections to upper limb motoneurones in humans. J Physiol (Lond) 448:397–412

Pessiglione M, Seymour B, Flandin G, Dolan RJ, Frith CD (2006) Dopamine-dependent prediction errors underpin reward-seeking behaviour in humans. Nature 442:1042–1045

Petrides M, Pandya DN (2009) Distinct parietal and temporal pathways to the homologues of Broca’s area in the monkey. PLoS Biol 7:e1000170

Pin JP, Duvoisin R (1995) The metabotropic glutamate receptors: structure and functions. Neuropharmacology 34:1–26

Pincze Z, Lakatos P, Rajkai C, Ulbert I, Karmos G (2002) Effect of deviant probability and interstimulus/interdeviant interval on the auditory N1 and mismatch negativity in the cat auditory cortex. Brain Res Cogn Brain Res 13:249–253

Pisani A, Martella G, Tscherter A, Bonsi P, Sharma N, Bernardi G, Standaert DG (2006) Altered responses to dopaminergic D2 receptor activation and N-type calcium currents in striatal cholinergic interneurons in a mouse model of DYT1 dystonia. Neurobiol Dis 24:318–325

Pons TP, Kaas JH (1986) Corticocortical connections of area 2 of somatosensory cortex in macaque monkeys: a correlative anatomical and electrophysiological study. J Comp Neurol 248:313–335

Porter R (1985) The corticomotoneuronal component of the pyramidal tract: corticomotoneuronal connections and functions in primates. Brain Res 357:1–26

Porter R, Hore J (1969) Time course of minimal corticomotoneuronal excitatory postsynaptic potentials in lumbar motoneurons of the monkey. J Neurophysiol 32:443–451

Prabhu G, Shimazu H, Cerri G, Brochier T, Spinks RL, Maier MA, Lemon RN (2009) Modulation of primary motor cortex outputs from ventral premotor cortex during visually guided grasp in the macaque monkey. J Physiol (Lond) 587:1057–1069

Preuss TM, Goldman-Rakic PS (1989) Connections of the ventral granular frontal cortex of macaques with perisylvian premotor and somatosensory areas: anatomical evidence for somatic representation in primate frontal association cortex. J Comp Neurol 282:293–316

Prevosto V, Graf W, Ugolini G (2010) Cerebellar inputs to intraparietal cortex areas LIP and MIP: functional frameworks for adaptive control of eye movements, reaching, and arm/eye/head movement coordination. Cereb Cortex 20:214–228

Pruszynski JA, Kurtzer I, Nashed JY, Omrani M, Brouwer B, Scott SH (2011) Primary motor cortex underlies multi-joint integration for fast feedback control. Nature 478:387–390

Przybyszewski AW, Gaska JP, Foote W, Pollen DA (2000) Striate cortex increases contrast gain of macaque LGN neurons. Vis Neurosci 17:485–494

Rao RP, Ballard DH (1999) Predictive coding in the visual cortex: a functional interpretation of some extra-classical receptive-field effects. Nat Neurosci 2:79–87

Raptis H, Burtet L, Forget R, Feldman AG (2010) Control of wrist position and muscle relaxation by shifting spatial frames of reference for motoneuronal recruitment: possible involvement of corticospinal pathways. J Physiol (Lond) 588:1551–1570

Rathelot J-A, Strick PL (2006) Muscle representation in the macaque motor cortex: an anatomical perspective. Proc Natl Acad Sci USA 103:8257–8262

Rathelot J-A, Strick PL (2009) Subdivisions of primary motor cortex based on cortico-motoneuronal cells. Proc Natl Acad Sci USA 106:918–923

Rauss K, Schwartz S, Pourtois G (2011) Top-down effects on early visual processing in humans: a predictive coding framework. Neurosci Biobehav Rev 35:1237–1253

Reichova I, Sherman SM (2004) Somatosensory corticothalamic projections: distinguishing drivers from modulators. J Neurophysiol 92:2185–2197

Rivadulla C, Sharma J, Sur M (2001) Specific roles of NMDA and AMPA receptors in direction-selective and spatial phase-selective responses in visual cortex. J Neurosci 21:1710–1719

Rockland KS, Drash GW (1996) Collateralized divergent feedback connections that target multiple cortical areas. J Comp Neurol 373:529–548

Rockland KS, Pandya DN (1979) Laminar origins and terminations of cortical connections of the occipital lobe in the rhesus monkey. Brain Res 179:3–20

Rodriguez-Oroz MC, Jahanshahi M, Krack P, Litvan I, Macias R, Bezard E, Obeso JA (2009) Initial clinical manifestations of Parkinson’s disease: features and pathophysiological mechanisms. Lancet Neurol 8:1128–1139

Rozzi S, Calzavara R, Belmalih A, Borra E, Gregoriou GG, Matelli M, Luppino G (2006) Cortical connections of the inferior parietal cortical convexity of the macaque monkey. Cereb Cortex 16:1389–1417

Rudolf GD, Cronin CA, Landwehrmeyer GB, Standaert DG, Penney JB, Young AB (1996) Expression of N-methyl-d-aspartate glutamate receptor subunits in the prefrontal cortex of the rat. Neuroscience 73:417–427

Rudomin P, Schmidt RF (1999) Presynaptic inhibition in the vertebrate spinal cord revisited. Exp Brain Res 129:1–37

Sáez I, Friedlander MJ (2009) Synaptic output of individual layer 4 neurons in guinea pig visual cortex. J Neurosci 29:4930–4944

Sainburg RL, Poizner H, Ghez C (1993) Loss of proprioception produces deficits in interjoint coordination. J Neurophysiol 70:2136–2147

Sainburg RL, Ghilardi MF, Poizner H, Ghez C (1995) Control of limb dynamics in normal subjects and patients without proprioception. J Neurophysiol 73:820–835

Salin PA, Bullier J (1995) Corticocortical connections in the visual system: structure and function. Physiol Rev 75:107–154

Salt TE (2002) Glutamate receptor functions in sensory relay in the thalamus. Philos Trans R Soc Lond B Biol Sci 357:1759–1766

Sandell JH, Schiller PH (1982) Effect of cooling area 18 on striate cortex cells in the squirrel monkey. J Neurophysiol 48:38–48

Schieber MH (2007) Comparative anatomy and physiology of the corticospinal system. Handb Clin Neurol 82:15–37

Schiller J, Major G, Koester HJ, Schiller Y (2000) NMDA spikes in basal dendrites of cortical pyramidal neurons. Nature 404:285–289

Scott S (2004) Optimal feedback control and the neural basis of volitional motor control. Nat Rev Neurosci 5:532–546

Schmidlin E, Brochier T, Maier MA, Kirkwood PA, Lemon RN (2008) Pronounced reduction of digit motor responses evoked from macaque ventral premotor cortex after reversible inactivation of the primary motor cortex hand area. J Neurosci 28:5772–5783

Self MW, Kooijmans RN, Supèr H, Lamme VA, Roelfsema PR (2012) Different glutamate receptors convey feedforward and recurrent processing in macaque V1. Proc Natl Acad Sci USA 109:11031–11036

Seki K, Fetz EE (2012) Gating of sensory input at spinal and cortical levels during preparation and execution of voluntary movement. J Neurosci 32:890–902

Shadmehr R, Smith MA, Krakauer JW (2010) Error correction, sensory prediction, and adaptation in motor control. Annu Rev Neurosci 33:89–108

Shapovalov AI (1975) Neuronal organization and synaptic mechanisms of supraspinal motor control in vertebrates. Rev Physiol Biochem Exp Pharmacol 72:1–54

Shemmell J, Krutky MA, Perreault EJ (2010) Stretch sensitive reflexes as an adaptive mechanism for maintaining limb stability. Clin Neurophysiol 121:1680–1689

Sherman SM, Guillery RW (1998) On the actions that one nerve cell can have on another: distinguishing “drivers” from “modulators”. Proc Natl Acad Sci USA 95:7121–7126

Shima K, Tanji J (1998) Involvement of NMDA and non-NMDA receptors in the neuronal responses of the primary motor cortex to input from the supplementary motor area and somatosensory cortex: studies of task-performing monkeys. Jpn J Physiol 48:275–290

Shimazu H, Maier MA, Cerri G, Kirkwood PA, Lemon RN (2004) Macaque ventral premotor cortex exerts powerful facilitation of motor cortex outputs to upper limb motoneurons. J Neurosci 24:1200–1211

Shinoda Y, Yokota J, Futami T (1981) Divergent projection of individual corticospinal axons to motoneurons of multiple muscles in the monkey. Neurosci Lett 23:7–12

Shipp S (2005) The importance of being agranular: a comparative account of visual and motor cortex. Philos Trans R Soc Lond B Biol Sci 360:797–814

Shipp S (2007) Structure and function of the cerebral cortex. Curr Biol 17:R443–R449

Shipp S, Zeki S (1989a) The organization of connections between areas V5 and V1 in macaque monkey visual cortex. Eur J Neurosci 1:309–332

Shipp S, Zeki S (1989b) The organization of connections between areas V5 and V2 in macaque monkey visual cortex. Eur J Neurosci 1:333–354

Shipp S, Blanton M, Zeki S (1998) A visuo-somatomotor pathway through superior parietal cortex in the macaque monkey: cortical connections of areas V6 and V6A. Eur J Neurosci 10:3171–3193

Shlosberg D, Amitai Y, Azouz R (2006) Time-dependent, layer-specific modulation of sensory responses mediated by neocortical layer 1. J Neurophysiol 96:3170–3182

Sloper JJ, Hiorns RW, Powell TP (1979) A qualitative and quantitative electron microscopic study of the neurons in the primate motor and somatic sensory cortices. Philos Trans R Soc Lond B Biol Sci 285:141–171

Spratling MW (2010) Predictive coding as a model of response properties in cortical area V1. J Neurosci 30:3531–3543

Stepniewska I, Preuss TM, Kaas JH (1993) Architectonics, somatotopic organization, and ipsilateral cortical connections of the primary motor area (M1) of owl monkeys. J Comp Neurol 330:238–271

Stepniewska I, Sakai ST, Qi H-X, Kaas JH (2003) Somatosensory input to the ventrolateral thalamic region in the macaque monkey: potential substrate for parkinsonian tremor. J Comp Neurol 455:378–395

Strick PL (2002) Stimulating research on motor cortex. Nat Neurosci 5:714–715

Summerfield C, Koechlin E (2008) A neural representation of prior information during perceptual inference. Neuron 59:336–347

Summerfield C, Trittschuh EH, Monti JM, Mesulam MM, Egner T (2008) Neural repetition suppression reflects fulfilled perceptual expectations. Nat Neurosci 11:1004–1006

Summerfield C, Wyart V, Johnen VM, de Gardelle V (2011) Human scalp electroencephalography reveals that repetition suppression varies with expectation. Front Hum Neurosci 5:67

Todorov E (2004) Optimality principles in sensorimotor control. Nat Neurosci 7:907–915

Todorov E, Jordan MI (2002) Optimal feedback control as a theory of motor coordination. Nat Neurosci 5:1226–1235

Tokuno H, Tanji J (1993) Input organization of distal and proximal forelimb areas in the monkey primary motor cortex: a retrograde double labeling study. J Comp Neurol 333:199–209

Tölle TR, Berthele A, Zieglgansberger W, Seeburg PH, Wisden W (1993) The differential expression of 16 NMDA and non-NMDA receptor subunits in the rat spinal cord and in periaqueductal gray. J Neurosci 13:5009–5028

Tracey DJ, Asanuma C, Jones EG, Porter R (1980) Thalamic relay to motor cortex: afferent pathways from brain stem, cerebellum, and spinal cord in monkeys. J Neurophysiol 44:532–554

Traynelis SF, Wollmuth LP, McBain CJ, Menniti FS, Vance KM, Ogden KK, Hansen KB, Yuan H, Myers SJ, Dingledine R (2010) Glutamate receptor ion channels: structure, regulation, and function. Pharmacol Rev 62:405–496

Turner JP, Salt TE (1998) Characterization of sensory and corticothalamic excitatory inputs to rat thalamocortical neurones in vitro. J Physiol (Lond) 510:829–843

Vallbo AB (1970) Discharge patterns in human muscle spindle afferents during isometric voluntary contractions. Acta Physiol Scand 80:552–566

Velliste M, Perel S, Spalding MC, Whitford AS, Schwartz AB (2008) Cortical control of a prosthetic arm for self-feeding. Nature 453:1098–1101

Viaene AN, Petrof I, Sherman SM (2011a) Properties of the thalamic projection from the posterior medial nucleus to primary and secondary somatosensory cortices in the mouse. Proc Natl Acad Sci USA 108:18156–18161

Viaene AN, Petrof I, Sherman SM (2011b) Synaptic properties of thalamic input to layers 2/3 and 4 of primary somatosensory and auditory cortices. J Neurophysiol 105:279–292

Viaene AN, Petrof I, Sherman SM (2011c) Synaptic properties of thalamic input to the subgranular layers of primary somatosensory and auditory cortices in the mouse. J Neurosci 31:12738–12747

Vitek JL, Ashe J, DeLong MR, Alexander GE (1994) Physiologic properties and somatotopic organization of the primate motor thalamus. J Neurophysiol 71:1498–1513

Vuust P, Ostergaard L, Pallesen KJ, Bailey C, Roepstorff A (2009) Predictive coding of music—brain responses to rhythmic incongruity. Cortex 45:80–92

Wall PD, Lidierth M (1997) Five sources of a dorsal root potential: their interactions and origins in the superficial dorsal horn. J Neurophysiol 78:860–871

Watanabe-Sawaguchi K, Kubota K, Arikuni T (1991) Cytoarchitecture and intrafrontal connections of the frontal cortex of the brain of the Hamadryas baboon (Papio hamadryas). J Comp Neurol 311:108–133

Witham CL, Wang M, Baker SN (2010) Corticomuscular coherence between motor cortex, somatosensory areas and forearm muscles in the monkey. Front Syst Neurosci 4:38

Wolpert DM, Kawato M (1998) Multiple paired forward and inverse models for motor control. Neural Netw 11:1317–1329

Wong YC, Kwan HC, Mackay WA, Murphy JT (1978) Spatial organization of precentral cortex in awake primates I. Somatosensory inputs. I. Somatosensory inputs. J Neurophysiol 41:1107–1119

Zeki S, Shipp S (1988) The functional logic of cortical connections. Nature 335:311–317

Ziemann U, Ilić TV, Alle H, Meintzschel F (2004) Estimated magnitude and interactions of cortico-motoneuronal and Ia afferent input to spinal motoneurones of the human hand. Neurosci Lett 364:48–52

Zilberter M, Holmgren C, Shemer I, Silberberg G, Grillner S, Harkany T, Zilberter Y (2009) Input specificity and dependence of spike timing-dependent plasticity on preceding postsynaptic activity at unitary connections between neocortical layer 2/3 pyramidal cells. Cereb Cortex 19:2308–2320

Zilles K, Schlaug G, Matelli M, Luppino G, Schleicher A, Qü M, Dabringhaus A, Seitz R, Roland PE (1995) Mapping of human and macaque sensorimotor areas by integrating architectonic, transmitter receptor. MRI and PET data. J Anat 187(Pt 3):515–537