Single-Neuron Theory of Consciousness

Sevush, Steven (2005) Single-Neuron Theory of Consciousness. [Preprint]

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By most accounts, the mind arises from the integrated activity of large populations of neurons distributed across multiple brain regions. A contrasting model is presented in the present paper that places the mind/brain interface not at the whole brain level but at the level of single neurons. Specifically, it is proposed that each neuron in the nervous system is independently conscious, with conscious content corresponding to the spatial pattern of a portion of that neuron's dendritic electrical activity. For most neurons, such as those in the hypothalamus or posterior sensory cortices, the conscious activity would be assumed to be simple and unable to directly affect the organism's macroscopic conscious behavior. For a subpopulation of layer 5 pyramidal neurons in the lateral prefrontal cortices, however, an arrangement is proposed to be present such that, at any given moment: i) the spatial pattern of electrical activity in a portion of the dendritic tree of each neuron in the subpopulation individually manifests a complexity and diversity sufficient to account for the complexity and diversity of conscious experience; ii) the dendritic trees of the neurons in the subpopulation all contain similar spatial electrical patterns; iii) the spatial electrical pattern in the dendritic tree of each neuron interacts nonlinearly with the remaining ambient dendritic electrical activity to determine the neuron's overall axonal response; iv) the dendritic spatial pattern is reexpressed at the population level by the spatial pattern exhibited by a synchronously firing subgroup of the conscious neurons, thereby providing a mechanism by which conscious activity at the neuronal level can influence overall behavior. The resulting scheme is one in which conscious behavior appears to be the product of a single macroscopic mind, but is actually the integrated output of a chorus of minds, each associated with a different neuron.

Item Type:Preprint
Keywords:binding problem; consciousness; hard problem; neural correlate of consciousness; single-neuron theory
Subjects:Philosophy > Philosophy of Mind
Neuroscience > Behavioral Neuroscience
Biology > Theoretical Biology
ID Code:4432
Deposited By:Sevush, Dr. Steven
Deposited On:06 Jul 2005
Last Modified:11 Mar 2011 08:56

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Select the SEEK icon to attempt to find the referenced article. If it does not appear to be in cogprints you will be forwarded to the paracite service. Poorly formated references will probably not work.

Abeles M, 1991: Corticonics: Neural Circuits of the Cerebral Cortex. New York: Cambridge University Press.

Abeles M, 2004: Time is precious. Science 304:523-524.

Adrian ED, 1926: J Physiol 61:47.

Aertsen A, Diesmann M, Gewaltig MO, 1996: Propagation of synchronous spiking activity in feedforward neural networks. J Physiol Paris 90:243-247.

Agmon-Snir H, Carr CE, Rinzel J, 1998: The role of dendrites in auditory coincidence detection. Nature 393:268-272.

Alexander GE, Delong MR, Strick PL, 1986: Parallel organization of functionally segregated circuits linking basal ganglia and cortex. Annu Rev Neurosci 9:357-381.

Alexander DM, Bourke PD, Sheridan P, Konstandatos O, Wright JJ, 2004: Intrinsic connections in tree shrew V1 imply a global to local mapping. Vision Research 44:857-876.

Alexander MP, Stuss DT, 2000: Disorders of frontal lobe functioning. Seminars in Neurology 20:427-437.

Amaral DG, Price JL, 1984: Amydalo-cortical projections in the monkey (Macaca fascicularis). J Comp Neurol 230:465-496.

Anderson JC, Binzegger T, Kahana O, Martin KAC, Segev I, 1999: Dendritic asymmetry cannot account for directional responses of neurons in visual cortex. Nature Neuroscience 2:820-824.

Aou S, Oomura Y, Hishino H, Ono T, Yambe K, Sikdar SK, Noda T, Inoue M, 1983: Functional heterogeneity of single neuronal activity in the monkey dorsolateral prefrontal cortex. Brain Res 260:121-124.

Arnoldi HM, Brauer W, 1996: Synchronization without oscillatory neurons. Biol Cybern 74:209-223.

Arnoldi HM, Englmeier KH, Brauer W, 1999: Translation-invariant pattern recognition based on Synfire chains. Biol Cybern 80:433-447.

Averbeck BB, Lee D, 2004: Coding and transmission of information by neural ensembles. Trends in Neurosciences 4:225-230.

Aviel Y, Pavlov E, Abeles M, Horn D, 2002: Synfire chain in a balanced network. Neurocomputing 44-46:285-292.

Baars BJ, 1988: A Cognitive Theory of Consciousness, Cambridge University Press.

Baars BJ, Franklin S, 2003: How conscious experience and working memory interact. Trends in Cognitive Sciences 7:166-172.

Baddeley A, 2003: Working memory: looking back and looking forward. Nature Reviews Neuroscience 4:829-839.

Baleydier C, Mauguiere F, 1980: The duality of the cingulated gyrus in monkey. Neuroanatomical study and functional hypothesis. Brain 103:525-554.

Barbas H, 1986: Pattern in the laminar origin of corticocortical connections. J Comp Neurol 252:415-422.

Barbas H, De Olmos J, 1990: Projections from the amygdala to basoventral and mediodorsal prefrontal regions in the rhesus monkey. J Comp Neurol 300:549-571.

Barbas H, Hilgetag CC, 2002: Rules relating connections to cortical structure in primate prefrontal cortex. Neurocomputing 44-46:301-308.

Barbas H, Pandya DN, 1989: Architecture and intrinsic connections of the prefrontal cortex in the rhesus monkey. J Comp Neurol 286:353-375.

Barbas H, Rempel-Clower N, 1997: Cortical structure predicts the pattern of corticocortical connections. Cerebral Cortex 7:635-646.

Barlow HB, 1972: Single units and sensation: a neuron doctrine for perceptual psychology? Perception 1:371-394.

Bates JF, Goldman-Rakic PS, 1993: Prefrontal connections of medial motor areas in the rhesus monkey. J Comp Neurol 336:211-228.

Baylis LL, Rolls ET, Baylis GC, 1995: Afferent connections of the caudolateral orbitofrontal cortex taste area of the primate. Neuroscience 64:801-812.

Bayne T, Chalmers DJ, 2003: What is the unity of consciousness? In: Cleeremans A (ed), The Unity of Consciousness: Binding, Integration, Dissociation. Oxford University Press: New York.

Bieberich E, 2002: Recurrent fractal neural networks: a strategy for the exchange of local and global information processing in the brain. BioSystems 66:145-164.

Binzegger T, Douglas RJ, Martin KAC, 2005: Axons in cat visual cortex are topologically self-similar. Cerebral Cortex 15:152-165.

Bialek W, Rieke F, de Ruyter van Steveninck RR, Warland D, 1991: Reading a neural code. Science 252: 1854-1857.

Bogen JE, 1995: On the neurophysiology of consciousness: Part II. Constraining the semantic problem. Consciousness and Cognition 4:137-158.

Borst A, Theunissen FE, 1999: Information theory and neural coding. Nature Neuroscience 2: 947-957.

Braver TS, Barch DM, Kelley WM, Buckner RL, Cohen NJ, Miezin FM, Snyder AZ, Ollinger JM, Akbudak E, Conturo TE, Petersen SE, 2001: Direct comparison of prefrontal cortex regions engaged by working and long-term memory tasks. Neuroimage 14:48-59.

Briggs F, Callaway EM, 2005: Laminar patterns of local excitatory input to layer 5 neurons in macaque primary visual cortex. Cerebral Cortex 15:479-488.

Broca P, 1861: Perte de la parole, remollissement chronique et destruction partielle du lobe anterieur gauche du cerveau. Bull Soc Anthropol 2:235-238.

Bruce C, Desimone R, Gross CG, 1981: Visual properties of neurons in a polysensory area in superior temporal sulcus of the macaque. J Neurophysiol 46:369-384.

Bush G, Luu P, Posner MI, 2000: Cognitive and emotional influences in anterior cingulated cortex. Trends in Cognitive Science 4:215-222.

Campbell SR, Wang DL, Jayaprakash C, 1999: Synchrony and desynchrony in integrate-and-fire oscillators. Neural Comput 11:1595-1619.

Cannon RC, Wheal HV, Turner DA, 1999: Dendrites of classes of hippocampal neurons differ in structural complexity and branching patterns. Journal of Comparative Neurology 413:619-633.

Cash S, Yuste R, 1999: Linear summation of excitatory inputs by CA1 pyramidal neurons. Neuron 22:383-394.

Chalmers D, 1996: The Conscious Mind. New York: Oxford University Press.

Chavis DA, Pandya DN, 1976: Further observations on cortico-frontal connections in the rhesus monkey. Brain Res 117:369-386.

Cohen JD, Perlstein WM, Braver TS, Nystrom LE, Noll DC

Conde F, Lund JS, Lewis DA, 1996: The hierarchical development of monkey visual cortical regions as revealed by the maturation of parvalbumin-immunoreactive neurons. Developmental Brain Research 96:261-276.

Constantinidis C, Franowicz, Goldman-Rakic PS, 2001: Coding specificity in cortical microcircuits: a multiple-electrode analysis of primate prefrontal cortex. The Journal of Neuroscience 21:3646-3655.

Constantinidis C, Goldman-Rakic PS, 2002: Correlated discharges among putative pyramidal neurons and interneurons in the primate prefrontal cortex. J Neurophysiol 88:3487-3497.

Crick F, Koch C, 1990a: Towards a neurobiological theory of consciousness. Seminars in Neuroscience 2:263-275.

Crick F, Koch C, 1990b: Some reflections on visual awareness. Cold Spring Harbor Symp Quant Biol 55:953-962.

Crick F, 1994: The Astonishing Hypothesis. Simon and Schuster: New York.

Crick F, Koch C, 1998: Consciousness and Neuroscience. Cerebral Cortex 8:97-107.

Dahaene S, Kerszberg M, Changeux JP, 1998: A neuronal model of a global workspace in effortful cognitive tasks. PNAS 95:14529-14534.

Damasio AR, 1998: Investigating the biology of consciousness. Phil Trans R Soc Lond B 353:1879-1882.

Damasio AR, 1999: The Feeling of What Happens. Harcourt Brace and Company: New York.

Dandy WE, 1946: The location of the conscious center in the brain – the corpus striatum. Bull Johns Hopkins Hosp 79:34-58.

Dantzker JI, Callaway EM, 2000: Laminar sources of synaptic input to cortical inhibitory interneurons and pyramidal neurons. Nature Neuroscience 3:701-707.

Deacon TW, 1992: Cortical connections of the inferior arcuate sulcus cortex in the macaque brain. Brain Research 573:8-26.

Dehaene S, Kerszberg M, Changeux JP, 1998: A neuronal model of a global workspace in effortful cognitive tasks. Proc Nat Acad Sci 95:14529-14534.

Dennett DC, 1991: Consciousness Explained. Boston: Little, Brown and Company.

D'Esposito M, Aguirre GK, Zarahn E, Ballard D, Shin RK, Lease J, 1998: Functional MRI studies of spatial and nonspatial working memory. Cognitive Brain Research 7:1-13.

Diesmann M, Gewaltig MO, Aertsen A, 1999: Stable propagation of synchronous spiking in cortical neural networks. Nature 402:529-533.

Douglas RF, Martin KAC, 2004: Neuronal circuits of the neocortex. Annu Rev Neurosci 27:419-451.

Durstewitz D, Seamans JK, Sejnowski TJ, 2000: Neurocomputational models of working memory. Nature Neuroscience Suppl 3: 1184-1191.

Edelman GM, 1993: Neural darwinism - selection and reentrant signaling in higher brain-function. Neuron 10:115-125.

Edelman GM and Tononi G, 2000: A Universe of Consciousness: How Matter Becomes Imagination. New York: Basic Books.

Edwards JCW, 2005: Is consciousness only a property of individual cells? J Cons Studies (in press).

Elston GN, 2003: Cortex, cognition and the cell: new insights into the pyramidal neuron and prefrontal function. Cerebral Cortex 13:1124-1138.

Engel AK, Fries P, Konig P, Brecht M, Singer W, 1999: Temporal binding, binocular rivalry, and consciousness. Consciousness and Cognition 8:128-151.

Eslinger PJ, Damasio AR, 1985: Severe disturbance of higher cognition after bilateral frontal lobe ablation: patient EVR. Neurology 35:1731-1741.

Felleman DJ, Van Essen DC, 1991: Distributed hierarchical processing in primate cerebral cortex. Cerebral Cortex 1:1-47.

Fletcher PC, Henson RNA, 2001: Frontal lobes and human memory: insights from functional neuroimaging. Brain 124:849-881.

Fries P, Reynolds JH, Rorie AE, DesimoneR, 2001: Modulation of oscillatory neuronal synchronization by selective visual attention. Science 291:1560-1563

Foxe JJ, Simpson GV, 2002: Flow of activation from V1 to frontal cortex in humans. A framework for defining "early" visual processing. Exp Brain Res 142:139-150.

Frick A, Magee J, Johnston D, 2004: LTP is accompanied by an enhanced local excitability of pyramidal neuron dendrites. Nat Neurosci 7:126-35

Froemke RC, Pou MM, Dan Y, 2005: Spike-timing-dependent synaptic plasticity depends on dendritic location. Nature 43:221-225.

Funahashi S, Inoue M, 2000: Neuronal interactions related to working memory processes in the primate prefrontal cortex revealed by cross-correlation analysis. Cerebral Cortex 10:533-551.

Fuster JM, 1998: Linkage at the top. Neuron 21:1223-1229.

Fuster JM, 1999: Memory in the cerebral cortex. MIT Press: Cambridge.

Fuster JM, Alexander GE, 1971: Neuron activity related to short-term memory. Science 173:652-654.

Geschwind N, 1981: The perverseness of the right hemisphere. The Behavioral and Brain Sciences 4:106-107.

Gewaltig MO, Diesmann M, Aertsen S, 2001: Propagation of cortical synfire activity: survival probability in single trials and stability in the mean. Neural Networks 14:657-673.

Gilbert CD, 1998: Adult cortical dynamics. Physiological Reviews 78:467-485.

Gilbert CD, Wiesel TN, 1986: Intrinsic connectivity and receptive field properties in visual cortex. Vision Res 25:365-374.

Gold I, 1999: Does 40-Hz oscillation play a role in visual consciousness? Consciousness and Cognition 8:186-195.

Golding NL, Kath WL, Spruston N, 2001: Dichotomy of action-potential backpropagation in CA1 pyramidal neuron dendrites. J Neurophysiol 10:2998-3010.

Goldman PS, Nauta WJ, 1976: Autoradiographic demonstration of a projection from prefrontal association cortex to the superior colliculus in the rhesus monkey. Brain Res 116:145-149.

Goldman-Rakic PS, 1992: Working memory and the mind. Scientific American September.

Goldman-Rakic PS, Schwartz ML, 1982: Interdigitation of contralateral and ipsilateral columnar projections to frontal association cortex in primates. Science 216:755-757.

Goldman-Rakic PS, Selemon LD, Schwartz ML, 1984: Dual pathways connecting the dorsolateral prefrontal cortex with the hippocampal formation and parahippocampal cortex in the rhesus monkey. Neuroscience 12:719-743.

Gonzalez-Burgos G, Barrionuevo G, Lewis DA, 2000: Horizontal synaptic connections in monkey prefrontal cortex: an in vitro electrophysiological study. 10:82-92.

Goodale MA, Milner AD, 1992: Separate visual pathways for perception and action. Trends Neurosci 15:20-25.

Grammont F, Riehle A, 2003: Spike synchronization and firing rate in a population of motor cortical neurons in relation to movement direction and reaction time. Biological Cybernetics 88:360-373.

Gray CM, 1999: The temporal correlation hypothesis of visual feature interaction: still alive and well. Neuron 24:31-47.

Gray JR, Braver TS, Raichle ME, 2002: Integration of emotion and cognition in the lateral prefrontal cortex. PNAS 99:4115-4120.

Grossberg S, Mingolla E, Pack C, 1999: A neural model of motion processing and visual navigation by cortical area MST. Cerebral Cortex 9:878-895.

Grush R, Churchland PS, 1995: Gaps in Penrose's toilings. J of Consciousness Studies 2:10-29.

Hardcastle V, 1994: Psychology's binding problem and possible neurobiological solutions. J of Consciousness Studies 1:66-90.

Hardcastle V, 1997: Consciousness and the neurobiology of perceptual binding. Seminars in Neurology 17:163-170.

Hausser M, 2001: Synaptic function: dendritic democracy. Curr Biol 11:R10-R12.

Hausser M, Major G, Stuart GJ, 2001: Differential shunting of EPSPs by action potentials. Science 291:138-141.

Hausser M, Mel B, 2003: Dendrites: bug or feature? Current Opinions in Neurobiology 13:372-383.

Heilman KM. and Valenstein EV, 1993: Clinical Neuropsychology. New York: Oxford University Press.

Helmchen F, Svoboda K, Denk W, Tank DW, 1999: In vivo dendritic calcium dynamics in deep-layer cortical pyramidal neurons. Nature Neuroscience 2:989-996.

Hertz J, Prugel-Bennett A, 1996: Learning synfire chains: turning noise into signal. Int J Neural Syst 7:445-450

Hoffman DA, Magee JC, Colbert CM, Johnston D: Potassium channel regulation of signal propagation in dendrites of hippocampal pyramidal neurons. Nature 38:869-875.

Hopfield JJ, 1982: Neural networks and physical systems with emergent collective computational abilities. PNAS 79:2554-2558.

Hopfield JJ, Herz AVM, 1995: Rapid local synchronization of action potentials: toward computation with coupled integrate-and-fire neurons. PNAS 92:6655-6662.

Hubel DH, Wiesel TN, 1962: Receptive fields, binocular interaction and functional architecture in the cat’s visual cortex. J Physiol (Lond) 160:106-154.

Hubel DH, Wiesel TN, 1965: Receptive fields and functional architecture in two non-striate areas (18 and 19) of the cat. J Neurophysiol 28:229-289.

Ikegaya Y, Aarn G, Cossart R, Aronov D, Lampl I, Ferster D, Yuste R: Synfire chains and cortical songs: temporal modules of cortical activity. Science 304:559-564.

Inui K, Wang X, Tamura Y, Kaneoke Y, Kakigi R, 2004: Serial processing in the human somatosensory system. Cerebral Cortex 14:851-857.

Ishai A, Ungerleider LG, Haxby JV, 2000: Distributed neural systems for the generation of visual images. Neuron 28:979-990.

Izhikevich EM, Gally JA, Edelman GM, 2004: Spike-timing dynamics of neuronal groups. Cerebral Cortex 14:933-944.

Jacobs GA, Theunissen FE, 2000: Extraction of sensory parameters from a neural map by primary sensory interneurons. The Journal of neuroscience 20:2934-2943.

James W, 1890: Principles of Psychology. Cambridge: Harvard University Press.

John ER, 2001: A field theory of consciousness. Consciousness and Cognition 10:184-213.

Johnson DMG, Illig KR, Behan M, Haberty LB, 2000: New features of connectivity in piriform cortex visualized by intracellular injection of pyramidal cells suggest that "primary" olfactory cortex functions like "association" cortex in other sensory systems. The Journal of Neuroscience 20:6974-6982.

Jones EG, 2002: Thalamic circuitry and thalamocortical synchrony. Phil Trans R Soc Lond B 357:1659-1673.

Jones EG, Powell TPS, 1970: An anatomical study of converging sensory pathways within the cerebral cortex of the monkey. Brain 93:793-820.

Koch C, 1999: Biophysics of Computation: Information Processing in Singe Neurons, Oxford University Press: Oxford.

Koch C, 1997: Computation and the single neuron. Nature 385:207-210.

Kjaer TW, Nowak M, Kjaer KW, Lou AR, Lou HC, 2001: Precuneus-prefrontal activity during awareness of visual verbal stimuli. Consciousness and Cognition 10:356-365.

Konig P, Engel AK, Roelfsema PR, Singer W, 1995: How precise is neural synchronization? Neural Computation 7:469-485.

Kritzer MF, Goldman-Rakic PS, 1995: Intrinsic circuit organization of the major layers and sublayers of the dorsolateral prefrontal cortex in the rhesus monkey. J Comp Neurol 359:131-143.

Kubota K, Niki H, 1971: Prefrontal cortical unit activity and delayed alternation performance in monkeys. J Neurophysiol 34:337-347.

Kuypers HGJM, Szwarcbart MK, Mishkin M, 1965: Occipitotemporal cortico-cortical connections in the rhesus monkey. Expl Neurol 11:245.

Lamme VAF, Roelfsema PR, 2000: The distinct modes of vision offered by feedforward and recurrent processing. Trends in Neurosciences 23:571-579.

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

Larkum ME, Kaiser KMM, Sakmann B, 1999b: Calcium electrogenesis in distal apical dendrites of layer 5 pyramidal cells at a critical frequency of back-propagating action potentials. Proc Natl Acad Sci USA 96:14600–14604.

Larkum ME, Zhu JJ, Sakmann B, 2001: Dendritic mechanisms underlying the coupling of the dendritic with the axonal action potential initiation zone of adult rat layer 5 pyramidal neurons. J Physiol 2001, 533:447-466.

Larkum ME, Senn W, Luscher HR, 2004: Top-down dendritic input increases the gain of layer 5 pyramidal neurons. Cerebral Cortex 14:1059-1070.

Larkum ME, Zhu JJ, 2002: Signaling of layer 1 and whisker-evoked Ca2+ and Na+ action potentials in distal and terminal dendrites of rat neocortical pyramidal neurons in vitro and in vivo. J Neurosci 22:6991-7005.

Lachaux JP, Rodriguez E, Martinerie J, Varela FJ, 1999: Measuring phase synchrony in brain signals. Hum Brain Mapp 8:194-208.

Laureys S, Lemaire C, Maquet P, 1999: Cerebral metabolism during vegetative state and after recovery to consciousness. J Neurol Neurosurg Psychiatry 67:121-133.

Laureys S, 2004: Functional neuroimaging in the vegetative state. NeuroRehabilitation 19:335-341.

Livingstone MS, 1998: Mechanisms of direction selectivity in macaque V1. Neuron 20:509-526.

Llinas R, Ribary U, Contreras D, Pedroarena C, 1998: The neuronal basis for consciousness. Phil Trans R Soc Lond B 353:1841-1849.

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.

Lubke J, Markram H, Frotscher M, Salmann B, 1996: Frequency and dendritic distribution of autapses established by layer 5 pyramidal neurons in the developing rat neocortex: comparison with synaptic innervation of adjacent neurons of the same class. J Neurosci 10:3209-3218.

Lumer ED, Edelman GM, Tononi G, 1997: Neural dynamics in a model of the thalamocortical system. II. The role of neural synchrony tested through perturbations of spike timing. Cereb Cortex 7:228-236.

MacGregor RJ, Ascarrunz FG, Kisley MA, 1995: Characterization, scaling, and partial representation of neural junctions and coordinated firing patterns by dynamic similarity. Biol Cybern 73:155-166.

Magee JC, 2000: Dendritic integration of excitatory synaptic input. Nature Reviews Neuroscience 1:181-190.

Magee JC, 1999: Dendritic Ih normalizes temporal summation in hippocampal CA1 neurons. Nat Neurosci 2:508-514

Magee JC, Cook EP, 2000: Somatic EPSP amplitude is independent of synapse location in hippocampal pyramidal neurons. Nature Neuroscience 3:895-903.

Mainen ZF, Sejnowski TJ, 1996: Influence of dendritic structure on firing pattern inn model neocortical neurons. Nature 382:363-366.

Marsalek P, Koch C, Maunsell J, 1997: On the relationship between synaptic input and spike output jitter in individual neurons. PNAS 94:735-740.

Maunsell JHR, Van Essen DC: The connections of the middle temporal visual area (MT) and their relationship to a cortical hierarchy in the macaque monkey. The Journal of Neuroscience 2563-2586.

McCullough W, Pitts W, 1943: A logical calculus of the ideas immanent in nervous activity. Bull Math Biophys 5:115-133.

McIntosh AR, Rajah MN, Lobaugh NJ, 1999: Interactions of prefrontal cortex in relation to awareness in sensory learning. Science 284:1531-1533.

Melchitzky DS, Gonzalez-Burgos G, Barrionuevo G, Lewis DA, 2001: Synaptic targets of the intrinsic axon collaterals of supragranular pyramidal neurons in monkey prefrontal cortex. J Comp Neurol 430:209-221.

Miller R, 1996: Cortico-thalamic interplay and the security of operation of neural assemblies and temporal chains in the cerebral cortex. Biol Cybern 75:263-275.

Miller EK, Cohen JD, 2001: An integrative theory of prefrontal cortex function. Annu Rev Neurosci 24:167-202.

Milner AD, Goodale MA, 1995: The Visual Brain in Action. Oxford University Press: Oxford.

Milner AD Goodale MA, 1993: Visual pathways to perception and action. Prog Brain Res 95:317-337.

Mitchell BD, Cauller LJ, 2001: Corticocortical and thalamocortical projections to layer 1 of the frontal neocortex in rats. Brain Res 921:68-77.

Mitchell BD, Macklis JD, 2005: Large-scale maintenance of dual projections by callosal and frontal cortical projection neurons in adult mice. J Comp Neurol 482:17-32.

Neltner L, Hansel D, Mato G, Meunier C, 2000: Synchrony in heterogeneous networks of spiking neurons. Neural Comput 12:1607-1641.

Niebur E, Hsiao SS, Johnson KO, 2002: Synchrony: a neuronal mechanism for attentional selection? Current Opinions in Neurobiology 12:190-194.

Niebur E, Koch C, 1994: A model for the neuronal implementation of selective visual attention based on temporal correlation among neurons. J Comput Neurosci 1:141-158.

Niebur E, Koch C, Rosin C, 1993: An oscillation-based model for the neural basis of attention. Vision Res 33:2789-2802.

Nirenberg S, Latham PE, 2003: Decoding neuronal spike trains: How important are correlations? PNAS 100:7348-7353.

Nowotny T, Huerta R, 2003: Explaining synchrony in feed-forward networks: are McCulloch-Pitts neurons good enough? Biol Cybern 89:237-241.

Oliveri M, Purriziani P, Carlesimo GA, Koch G, Tomaiuolo F, Panella M, Caltagirone C, 2001: Parieto-frontal interactions in visual-object and visual-spatial working memory: evidence from transcranial magnetic stimulation. Cerebral Cortex 11:606-618.

Orpwood RD, 1994: A possible neural mechanism underlying consciousness based on the pattern processing capabilities of pyramidal neurons in the cerebral cortex. J Theor Biol 169:403-418.

Pandya DN, Kuypers HG, 1969: Cortico-cortical connections in the rhesus monkey. Brain Research 13:13-48.

Pandya DN, Yeterian EH, 1990: Prefrontal cortex in relation to other cortical areas in rhesus monkey - architecture and connections. Prog Brain Res 85:63-94.

Parvizi J, Damasio A, 2001: Consciousness and the brainstem. Cognition 49:135-159.

Petrides M, Pandya DN, 1984: projections to the frontal cortex from the posterior parietal region in the rhesus monkey. J Comp Neurol 228:106-116.

Petrides M, Pandya DN, 1999: Dorsolateral prefrontal cortex: comparative cytoarchitectonic analysis in the human and the macaque brain and corticocortical connection pattern. Eur J Neurosci 11:1011-1036.

Phan KL, Wager T, Taylor SF, Liberzon I, 2002: Functional neuroanatomy of emotion: a meta-analysis of emotion activation studies in PET and fMRI. Neuroimage 16:331-348.

Pinsky PF, Rinzel J, 1995: Synchrony measures for biological neural networks. Biol Cybern 73:129-137.

Poirzai P, BrannonT, Mel BW, 2003: Pyramidal neuron as two-layer neural network. Neuron 37:989-999.

Polsky A, Mel BW, Schiller J, 2004: Computational subunits in thins dendrites of pyramidal cells. Nature Neuroscience 7:621-627.

Porrino LJ, Crane AM, Goldman-Rakic PS, 1981: Direct and indirect pathways from the amygdala to the frontal lobe in rhesus monkeys. J Comp Neurol 198:121-136.

Porter R, Ghosh S, Lange GD, Smith TG, 1991: A fractal analysis of pyramidal neurons in mammalian motor cortex. Neuroscience letters 130:112-116.

Postma EO, van den Herik HJ, Hudson PT, 2996: Robust feedforward processing in synfire chains. Int J Neural Syst 7:537-542.

Pouget A, Latham P, 2002: Digitalized neural networks: long-term stability from forgetful neurons. Nature Neuroscience 5:709-710.

Prut Y, Vaadia E, Bergman H, Haalman I, Slovin H, Abeles M, 1998: Spatiotemporal structure of cortical activity: properties and behavioral relevance. J Neurophysiol 79:2857-2874.

Puccetti R, 1973: Brain bisection and personal identity. British Journal of Philosophy of Science 24:339-355.

Raichle ME, 2000: The neural correlates of consciousness: an analysis of cognitive skill learning. In Gazzaniga MS: The New Cognitive Neurosciences, Second Edition, MIT Press: Cambridge.

Rainer G, Asaad WF, Miller EK, 1998: Memory fields of neurons in the primate prefrontal cortex. PNAS 95:15008-15013.

Rainer G, Rao SC, Miller EK, 1999: Prospective coding for objects in the primate prefrontal cortex. J Neurosci 19:5493-5505.

Rall W, 1959: Branching dendritic trees and motoneuron membrane resistivity. Exp Neurol 1:491-527.

Rall W, 1967: Distinguishing theoretical synaptic potentials computed for different soma-dendritic distributions of synaptic input. Journal of Neurophysiology 30:1138-1168.

Rammont F, Riehle A, 2003: Spike synchronization and firing rate in a population of motor cortical neurons in relation to movement direction and reaction time. Biol Cybernetics 88:30-373.

Rao SC, Rainer G, Miller EK, 1997: Integration of what and where in the primate prefrontal cortex. Science 276:821-824.

Rempel-Clower NL, Barbas H, 2000: The laminar pattern of connections between prefrontal and anterior temporal cortices in the rhesus monkey is related to cortical structure and function. Cerebral Cortex 10:851-865.

Revonsuo A, 1999: Binding and the phenomenal unity of consciousness. Consciousness and Cognition 8:178-185.

Reyes AD, 2003: Synchrony-dependent propagation of firing rate in iteratively constructed networks in vitro. Nature Neurosci 6: 593-599.

Reynolds JH, Desimone R, 1999: The role of neural mechanisms of attention in solving the binding problem Neuron 24:19-29.

Robertson LC, 2003: Binding, spatial attention and perceptual awareness. Nature Reviews Neuroscience 4:93-102.

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

Rolls ET, France L, Aggelopoulos NC, Reece S, 2003: An information theoretic approach to the contributions of the firing rates and the correlations between the firing of neurons. J Neurophysiol 89:2810-2822.

Romo R, Brody CD, Hernandez A, Lemus L, 1999: Neuronal correlates of parametric working memory in the prefrontal cortex. Nature 399:470-473.

Rudd ME, Brown LG, 1997: Noise adaptation in integrate-and-fire neurons. Neural Comput 9:1047-1069.

Sato M, Kuroda R, Ioku M, Kim A, Tanaka S, Nakakita K, Kohama A, Fugii C, Ono S, 1989: Regional cerebral blood flow in the resistant vegetative state. Neurol Med Chir 29:389-394.

Sauve K, 1999: Gamma-band synchronous oscillations: recent evidence regarding their functional significance. Consciousness and Cognition 8:213-224.

Schaefer AT, Larkum ME, Sakmann B, Roth A, 2003: Coincidence detection in pyramidal neurons is tuned by their dendritic branching pattern. J Neurophysiol 89:3143–3154.

Schiller J, Helmchen F, Sakmann B, 1997: Calcium action potentials restricted to distal apical dendrites of rat neocortical pyramidal neurons. J Physiol 505:605-616.

Schiller J, Schiller Y, Stuart G, Sakmann B, 1997: Calcium action potentials restricted to distal apical dendrites of rat neocortical pyramidal neurons. J Physiol 505, 605–616.

Schmahmann JD, Pandya DN, 1997: Anatomic organization of the basilar pontine projections from prefrontal cortices in rhesus monkey. J Neurosci 17:438-458.

Schubert D, Staiger JF, Cho N, Kotter R, Zilles K, Luhmann HJ, 2001: Layer-specific intracolumnar and transcolumnar functional connectivity of layer V pyramidal cells in rat barrel cortex. The Journal of Neuroscience 21:3580-3592.

Schuz A, 1998: Neuroanatomy in a computational perspective. In Arbib MA: Handbook of Brain Theory and neural Networks. MIT Press: Cambridge.

Schwartz EL, 1980: Computational anatomy and functional architecture of striate cortex: a spatial mapping approach to perceptual coding. Vision Research 20:645-669.

Schwindt P, Crill W, 1999: Mechanisms underlying burst and regular spiking evoked by

dendritic depolarization in layer 5 cortical pyramidal neurons. J Neurophysiol 81:1341–1354.

Seltzer B, Pandya DN, 1989: Frontal lobe connections of the superior temporal sulcus in rhesus monkey. J Comp Neurol 281:97-1113.

Shadlen MN, Newsome WT, 1994: Noise, neural codes and cortical organization. Curr Opin Neurobiol 4:569-579.

Shadlen MN, Newsome WT, 1998: The variable discharge of cortical neurons: implications for connectivity, computation, and information coding. J Neurosci 18:3870-3896.

Shadlen MN, Movshon JA, 1999: Synchrony unbound: a critical evaluation of the temporal binding hypothesis. Neuron 24:67-77.

Seager W, 1995: Consciousness, information and panpsychism. Journal of Consciousness Studies 2:272-288.

Segev I, London M, 2000: Untangling dendrites with quantitative models. Science 290:744-750.

Segev I, 2003: Synchrony is stubborn in feedforward cortical networks. Nature Neurosci 6:543-544.

Sewards TV, Sewards MA, 2001: On the correlation between synchronized oscillatory activities and consciousness. Consciousness and Cognition 10:485-495.

Singer W, 1998: Consciousness and the structure of neuronal representations. Phil Trans R Soc Lond B 353:1829-1840.

Singer W, 2001: Consciousness and the binding problem. Ann NY Acad Sci 929:123-146.

Singer W, Gray CM, 1995: Visual feature integration and the temporal correlation hypothesis. Annu Rev Neurosci 18:555-586.

Slotnick SD, Moo LR, Segal JB, Hart J, 1969: Distinct prefrontal cortex activity associated with item memory and source memory for visual shapes. Cognitive Brain Research 17:75-82.

Softky WR, Koch C, 1993: The highly irregular firing of cortical cells is inconsistent with temporal integration of random EPSPs. The Journal of Neuroscience 13:334-350.

Sougné JP, French RM, 2001: Synfire chains and catastrophic interference. In JD Moore and K Stenning (Eds.) Proceedings of the Twenty-Third Annual Conference of the Cognitive Science Society. Pp. 970-975. Mahwah,NJ: Lawrence Erbaum Ass.

Sperry RW, 1969: A modified concept of consciousness. Psychological Reviews 26:532-536.

Spratling MW, 2002: Cortical region interactions and the functional role of apical dendrites. Behavioral and Cognitive Neuroscience Reviews 1:219-228.

Spruston N, Schiller Y, Stuart G, Sakmann B, 1995: Activity-dependent action potential invasion and calcium influx into hippocampal CA1 dendrites. Science 286:297-300.

Spruston N, 2000: Distant synapses raise their voices. Nature Neuroscience 3:849-895.

Steinmetz PN, Roy A, Fitzgerald PJ, Hsiao SS, Johnson KO, Niebur E, 2000: Attention modulates synchronized neuronal firing in primate somatosensory cortex. Nature 404:187-190.

Stephan KM, Thaut MH, Wunderlich G, Schicks W, Tian B, Tellmann L, Schmitz T, Herzog H, McIntosh C, Seitz RJ, Homberg V, 2002: Conscious and subconscious sensorimotor synchronization - prefrontal cortex and the influence of awareness. Neuroimage 15:345-352.

Stevens CF, Zador AM, 1998: Input synchrony and the irregular firing of cortical neurons. Nature Neuroscience 1:210-217.

Stuart GJ, Sakmann B, 1994: Active propagation of somatic action potentials

into neocortical pyramidal cell dendrites. Nature 367:69–72.

Tanila H, Carlson S, Linnankoski I, Lindroos F, Kahila H, 1992: Functional properties of dorsolateral prefrontal cortical neurons in awake monkey. Behav Brain Res 47:169-180.

Taylor JG, 2001: The central role of the parietal lobes in consciousness. Consciousness and Cognition 10:379-417.

Thomson AM, Bannister AP, 1998: Postsynaptic pyramidal target selection by descending layer III pyramidal axons: dual intracellular recordings and biocytin filling in slices of rat neocortex. Neuroscience 3:669-683.

Thomson AM, Bannister AP, Mercer A, Morris OT, 2002a: Target and temporal pattern selection at neocortical synapses. Phil Trans R Soc Lond B 357:1781-1791.

Thomson AM, West DC, Wang Y, Bannister AP, 2002b: Synaptic connections and small circuits involving excitatory and inhibitory neurons in layers 22-5 of adult rat and cat neocrotex: triple intracellular recordings and biocytin labelling in vitro. Cerebral Cortex 12:936-953.

Thomson AM, Bannister AP, 2003: Interlaminar connections in the neocortex. Cerebral Cortex 13:5-17.

Thorpe S, 1998: Localized versus distributed representations. In: Arbib MA: Handbook of Brain Theory and Neural Networks. MIT Press: Boston.

Tulving E, 2002: Episodic memory: from mind to brain. Annu Rev Psychol. 2002;53:1-25.

Ungerleider LG, Mishkin M, 1982: Two cortical visual systems. In Analysis of Visual Behavior, DJ Ingle, MA Goodale, RJW Mansfield, eds. (Cambridge, MA: MIT Press).

Usrey WM, Reid RC, 1999: Synchronous activity in the visual system. Annu Rev Physiol 61:435-456.

Van Essen DC, Gallant JL, 1994: Neural mechanisms and motion processing in primate visual system. Neuron 13:1-10.

Van Ooyen A, Duijnhouwer J, Remme MWH, van Pelt J, 2002: The effect of dendritic topology on firing patterns in model neurons. Network Computation iin Neural Systems 13: 311-325.

Vetter P, Roth A, Hausser M, 2001: Propagation of action potentials in dendrites depends on dendritic morphology. J Neurophysiol 85:926-937.

Wei DS, Mei YA, Bagal A, Kao JP, Thompson SM, Tang CM, 2001: Compartmentalized and binary behavior of terminal dendrites in hippocampal pyramidal neurons. Science 293:2272-2275.

Weinberg RJ, Pierce JP, Rustioni A, 1990: Single fiber studies of ascending input to the cuneate nucleus of cats: I. Morphometry of primary afferent fibers. J Comp Neurol 300:113-133.

Wheeler MA, Stuss DT, Tulving E, 1995: Frontal lobe damage produces episodic memory impairment. Journal of the International Neuropsychological Society 1:525-536.

Whitehead AN, 1933: Adventures of Ideas. New York: The Free Press.

Williams SR, Stuart GJ, 2000: Backpropagation of physiological spike trains. J Neurosci 20:8238-8246.

Williams SR, Stuart GJ, 2002: Dependence of EPSP efficacy on synapse location in neocortical pyramidal neurons. Science 295:1907-1910.

Wolfe JM, Cave KR, 1999: The psychophysical evidence for a binding problem in human vision. Neuron 24:11-17.

Yazdanbakhsh A, Babadi B, Rouhani S, Arabzadeh E, Abbassian A, 2002: New attractor states for synchronous activity in synfire chains with excitatory and inhibitory coupling. Biol Cybern 86:367-378.

Zeki S, Bartels A, 1999: Toward a theory of visual consciousness. Consciousness and Cognition 8:225-259.


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