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framework and set it out under ten headings. This framework offers a coherent scheme for
explaining the neural correlates of (visual) consciousness in terms of competing cellular
assemblies. Most of the ideas we favor have been suggested before, but their combination is
original. We also outline some general experimental approaches to the problem and, finally,
acknowledge some relevant aspects of the brain that have been left out of the proposed
framework.
General strategy framework may well apply, however, to see, for example, either introns or RNA
The most difficult aspect of consciousness is other sensory modalities. We have been editing. And who would have guessed
the so-called hard problem of qualia1,2 especially interested in the alert macaque that DNA replication usually starts with
the redness of red, the painfulness of pain, monkey because to find the NCC, not the synthesis of a short stretch of RNA,
and so on. No one has produced any plau- only widespread neural activities but also which is then removed and replaced by
sible explanation as to how the experience the detailed behavior of single neurons (or DNA? The broad framework acted as a
of the redness of red could arise from the small groups of neurons) must be inves- guide, but careful experimentation was
actions of the brain. It appears fruitless to tigated on very fast time scales. This is dif- needed for the true details to be discov-
approach this problem head-on. Instead, ficult to do systematically in humans. ered. This lesson is broadly applicable
we are attempting to find the neural corre- Methods such as fMRI are too coarse in throughout biology.
late(s) of consciousness (NCC), in the hope both space and time to be of much use for
that when we can explain the NCC in causal our problem. On the other hand, experi- A preamble on the cerebral cortex
terms, this will make the problem of qualia ments in visual psychology are much eas- By the cortical system, we mean the cere-
clearer3. In round terms, the NCC is the ier to do on humans than on monkeys. bral cortex plus other regions closely asso-
minimal set of neuronal events that gives Moreover, humans can report what they ciated with it, such as the thalamus and
rise to a specific aspect of a conscious per- are conscious of and thus convey the con- the claustrum, and probably the basal
cept. We discuss elsewhere4 why we think tents of their consciousness. For these ganglia, the cerebellum and the many
consciousness has to be largely private. By reasons, experiments on monkeys and widespread brainstem projection systems.
private, we mean that it is accessible exclu- humans should be pursued in parallel. We shall refer to the front of the cor-
sively to the owner of the brain; it is impos- tex and the back of the cortex because
sible for me to convey to you the exact Framework the terms frontal and prefrontal can be
nature of my conscious percept of the color A framework is not a detailed hypothesis ambiguous. For example, is the anterior
red, though I can convey information about or set of hypotheses; rather, it is a sug- cingulate prefrontal?
it, such as whether two shades of red appear gested point of view for an attack on a sci- The dividing line between front and
to me to be the same or different. entific problem, often suggesting testable back is somewhat arbitrary. It roughly
Our main interest is not the enabling hypotheses. Biological frameworks differ coincides with the central sulcus. It may
factors needed for all forms of conscious- from frameworks in physics and chem- turn out that a good operational defini-
ness, such as the activity of the ascending istry because of the nature of evolution. tion is that the front is all those parts that
reticular systems in the brainstem. Rather, Biological systems do not have rigid laws, receive a significant input, via the thala-
we are interested in the general nature of as physics has. Evolution produces mech- mus, from the basal ganglia. (This simple
the neural activities that produce each anisms, and often sub-mechanisms, so division is probably not useful for olfac-
particular aspect of consciousness, such that there are few rules in biology which tion, however.)
as perceiving the specific color, shape or do not have occasional exceptions. There is an absolutely astonishing vari-
movement of an object. A good framework is one that sounds ety and specificity of actions performed
As a matter of tactics, we have con- reasonably plausible relative to available by the cortical system. The visual system
centrated on the visual system of primates, scientific data and that turns out to be of the higher mammals handles an almost
leaving on one side some of the more dif- largely correct. It is unlikely to be correct infinite variety of visual inputs and reacts
ficult aspects of consciousness, such as in all the details. A framework often con- to them in detail and with remarkable
emotion and self-consciousness. Our tains unstated (and often unrecognized) accuracy. It is clear that the system is high-
assumptions, but this is unavoidable. ly evolved, is likely to be specified epige-
Francis Crick is at the Salk Institute for An example from molecular biology netically in considerable detail and can
Biological Studies, 10010 N. Torrey Pines Road, might be helpful. The double-helical learn a large amount from experience.
La Jolla, California 92037, USA. structure of DNA immediately suggest- The main function of the sensory cor-
Christof Koch is at the California Institute of ed, in a novel way, the general nature of tex is to construct and use highly specific
Technology, 1200 East California Boulevard, gene composition, gene replication and feature detectors, such as those for orien-
Pasadena, California 91125, USA. gene action. This framework turned out tation, motion or faces. The features to
e-mail: koch@klab.caltech.edu to be broadly correct, but it did not fore- which any cortical neuron responds
are usually highly specific but multi- rather than following just one method (Further discussion of this idea comes at
dimensional. That is, one neuron does not very accurately. This appears to be how, the end of subhead #3 below.)
respond to a single feature but to a family for example, people see in depth. We have discussed elsewhere17 whether
of related features. Such features are some- Incoming visual information is often the neural activity in the front of the brain
times called the receptive field of that incomplete or ambiguous. If two similar is largely unconscious. One proposal18,19,
neuronthe non-classical receptive stimuli are presented in rapid succession, for example, is that humans are not direct-
field5 expresses the relevant context of the the brain blends them together into one ly conscious of their thoughts, but only of
classical receptive field. The visual fields percept. If they are different but in con- sensory representations of them in their
2003 Nature Publishing Group http://www.nature.com/natureneuroscience
of neurons higher in the visual hierarchy tradiction, such as a face and a house, the imagination. At the moment, there is no
are larger and respond to more complex brain selects one at a time (as in binocular consensus about this20.
features than those that are lower down. rivalry) instead of blending them togeth- The hypothesis of the homunculus
An important but neglected aspect of er. In cases where there is not enough is very much out of fashion these days,
the firing of a neuron (or a small group of information to lead to an unambiguous but this is, after all, how ever yone
associated neurons) is its projective interpretation of ones environment9, the thinks of themselves. It would be sur-
field6. This term describes the perceptual cortical networks fill inthat is, they prising if this overwhelming illusion
and behavioral consequences of stimulat- make their best guess, given the incom- did not reflect in some way the general
ing such a neuron in an appropriate man- plete information. Such filling-in is like- organization of the brain.
ner (for further discussion of motor and ly to happen in many places in the brain.
premotor cortex, see ref. 7). Both the This general principle is an important 2. Zombie modes and consciousness
receptive field and the projective field are guide to much of human behavior (as in Many actions in response to sensory
dynamic, not merely static, and both can jumping to conclusions). inputs are rapid, transient, stereotyped
be modified by experience. and unconscious 14,21 . They could be
How are feature detectors formed? A Our present framework thought of as cortical reflexes. Con-
broad answer is that neurons do this by Having outlined a few general points sciousness deals more slowly with broad-
detecting common and significant corre- about the cortical system, let us now con- er, less stereotyped aspects of the sensory
lations in their inputs and by altering their sider specifically the NCC and its atten- inputs (or a reflection of these in imagery)
synapses (and perhaps other properties) dant properties. We are mainly interested and takes time to decide on appropriate
so that they can more easily respond to in time periods on the order of a few hun- thoughts and responses. It is needed
such inputs. In other words, the brain is dred milliseconds, or at the most, several because otherwise, a vast number of dif-
very good at detecting apparent causation. seconds, so we can now leave on one side ferent zombie modes would be required.
Exactly how it does this is more contro- processes that take more time, such as The conscious system may interfere some-
versial. The main mechanism is probably permanently laying down a new memo- what with the concurrent zombie system.
Hebbian, but Hebbs seminal suggestion ry. We have listed the main ingredients of It seems to be a great evolutionary advan-
needs to be expanded. our framework under ten headings. We tage to have zombie modes that respond
All the above might suggest that cor- have not previously discussed in print rapidly, in a stereotyped manner, togeth-
tical action is highly local. Nothing could items 3, 5, 7 and 10, though we have men- er with a slightly slower system that allows
be further from the truth. In the cortex, tioned the first three in a book chapter time for thinking and planning more
there is continual and extensive interac- that is still in press10. Many of our basic complex behavior.
tion, both among neighboring cells and ideas on consciousness, such as the Visual zombie modes in the cortex
also very widely, thanks to the many long importance of attention and correlated probably use the dorsal stream in the pari-
cortico-cortical and cortico-thalamo-cor- firing, were outlined in our 1990 paper11, etal region14. Some parietal activity, how-
tical routes. This is much less true of the and in 1995, we suggested a plausible ever, also affects consciousness by
thalamus itself. function for consciousness 12 and later producing attentional effects on the ven-
The sensory cortex is arranged in a updated our ideas in a 1998 review3. Pre- tral stream, at least under some circum-
semi-hierarchical manner. (This is cer- viously we proposed that people are not stances. The conscious mode for vision
tainly true of the visual cortex8, but is less directly conscious of the neural activity in depends largely on the early visual areas
clear in the front of the brain.) That is, primary visual cortex (V1) and that the (beyond V1) and especially on the ventral
most cortical areas do not detect simple (visual) cortex appears hierarchical stream. There are no recorded cases of
correlations in the sensory input, but because it contains no strong loops12,13. purely parietal damage which led to a
detect correlations among correlations More recently, we have supported the sug- complete loss of conscious vision.
being expressed by other cortical areas. gestion14 that in addition to a slower, all- In a zombie mode, the main flow of
This remarkable feature of the cortex is purpose conscious mode, the brain has information is probably feed-forward. It
seldom emphasized. many zombie modes15, which are char- could be considered a forward-traveling
There has been a great selective advan- acterized by rapid and somewhat stereo- net-wave. A net-wave is a propagating
tage in reacting very rapidly, for both typed responses. wave of neural activity, but it is not the
predators and prey. For this reason, the same as a wave in a continuous medium.
best is the enemy of the good. Typically, it 1. The (unconscious?) homunculus Neural networks in a cortex have both
is better to achieve a rapid but occasion- A good way to begin to consider the overall short and long connections, so a net-wave
ally imperfect performance instead of a behavior of the cerebral cortex is to imag- may, in some cases, jump over interven-
more prolonged one that always produces ine that that the front of the brain is look- ing regions. In the conscious mode, it
a perfect result. Another general principle ing at the sensory systems, most of which seems likely that the flow is in both direc-
may be to use several rough and ready are at the back of the brain. This division of tions (see #5 below) so that it resembles
methods in parallel to reach a conclusion, labor does not lead to an infinite regress16. more of a standing net-wave10.
3. Coalitions of neurons If there are coalitions in the front of conscious unless there is an essential node
The cortex is a very highly and specifically the cortex, they may have a somewhat dif- for it. For consciousness, there may be
interconnected neural network. It has many ferent character from those formed at the other necessary conditions, such as pro-
types of excitatory and inhibitory interneu- back of the cortex. There may be more jecting to the front of the brain12.
rons and acts by forming transient coali- than one coalition that achieves winning A node, all by itself, cannot produce
tions of neurons, the members of which status and hence produces conscious consciousness. Even if the neurons in that
support one another. Coalitions implies experience. The coalitions in the front node were firing appropriately, this would
assembliesan idea which goes back at may reflect feelings such as happiness and, produce little effect if their output synaps-
2003 Nature Publishing Group http://www.nature.com/natureneuroscience
least to Hebb22plus competition among perhaps, the feeling of authorship, which es were inactivated. A node is a node, not
them (see also ref. 23). On the basis of is related to free will25. Such feelings may a network. Thus a particular coalition is
experimental results in the macaque, some be more diffuse and persist for a longer an active network, consisting of the rele-
researchers suggest that selective attention time than coalitions in the back of cortex. vant set of interacting nodes that tem-
biases the competition among rivalrous cell The terms affect and valuations are now porarily sustains itself.
assemblies, but they do not explicitly relate being used for what we have traditional- Much useful information can be
this idea to consciousness24. ly called feelings18,19. Our first working obtained from lesions. In humans, the
The various neurons in a coalition in assumption (the homunculus) implies damaged area is usually fairly large. It is
some sense support one another, either that it is better not to regard the back plus not clear what effects a very small, (possi-
directly or indirectly, by increasing the the front as one single coalition, but rather bly bilateral) reversible lesion would have
activity of their fellow members. The as two or more separate coalitions that in the macaque, as it is difficult to discov-
dynamics of coalitions are not simple. In interact massively, but not in an exactly er exactly what a monkey is conscious of.
general, at any moment the winning coali- reciprocal manner. The smallest useful node may be a corti-
tion is somewhat sustained, and embodies cal column29 or, perhaps, a portion of a
what we are conscious of. 4. Explicit representations cortical column. The feature which that
It may help to make a crude political An explicit representation of a particular node represents is (broadly) its columnar
analogy. The primaries and the early aspect of the visual scene implies that a property. This is because although a sin-
events in an election would correspond small set of neurons exists that responds gle type of pyramidal cell usually sends its
roughly to the preliminary unconscious as a detector for that feature, without fur- information to only one or two cortical
processing. The winning coalition associ- ther complex neural processing. A possi- areas, the pyramidal cells in a column pro-
ated with an object or event would corre- ble probe, or an operational test, for an ject the columnar property collectively to
spond to the winning party, which would explicit representation might be whether a many cortical areas, and can thus strength-
remain in power for some time and would single layer of neurons could deliver the en any coalition that is forming.
attempt to influence and control future correct answer. For example, if a single-
events. Attention would correspond to the layered neural network were fed the activ- 5. The higher levels first
efforts of journalists, pollsters and others ity of retinal neurons, it would not be able For a new visual input, the neural activity
to focus on certain issues rather than oth- to recognize a face. Fed from the relevant first travels rapidly and unconsciously up
ers, and thus attempt to bias the electorate parts of inferior temporal cortex, howev- the visual hierarchy to a high level, possi-
in their favor. Perhaps those large pyrami- er, it could reliably signal face or no face. bly in the front of the brain (this might
dal cells in cortical layer 5 that project to There is much evidence from both instantiate a zombie mode). Signals then
the superior colliculus and the thalamus humans and monkeys that if there are no start to move backward down the hierar-
(both involved in attention) would corre- such explicit neurons, or if they are all lost chy so that the first stages to reach con-
spond to electoral polls. These progress by brain damage, then the subject is sciousness are at the higher levels
from early, tentative polls to later, rather unable to consciously perceive that aspect (showing the gist of the scene30,31; see also
more accurate ones as the election directly. Well-known clinical examples are ref. 32), which send these conscious sig-
approaches. It is unlikely that all this hap- achromatopsia (loss of color perception), nals again to prefrontal cortex, followed
pens in the brain in a fixed time sequence. prosopagnosia (loss of face recognition) by corresponding activity at successive
The brain may resemble more the British or akinetopsia (loss of motion percep- lower levels (to provide the visual details).
system, in which the time between one tion). In all cases, one or a few attributes This is an oversimplified description.
election and the next can be irregular. Such of conscious experience have been lost, There are also many horizontal connec-
an analogy should not be pressed too far. while most other aspects remain intact. In tions in the hierarchy.
Like all analogies, it should be regarded as the macaque, a small, irreversible lesion How far up the hierarchy the initial
a possible source of ideas, which of course of the motion area MT/V5 leads to a tem- net-wave travels may depend upon
will have to be confirmed by experiment. poral deficit in motion perception that whether attention is diffused or focused
Coalitions can vary both in size and in recovers within days. Larger lesions cause at some particular level.
character. For example, a coalition pro- a more permanent loss26.
duced by visual imagination (with ones Note that an explicit representation is 6. Driving and modulating connections
eyes closed) may be less widespread than a a necessary but not sufficient condition In considering the physiology of coali-
coalition produced by a vivid and sus- for the NCC to occur. tions, it is especially important to under-
tained visual input from the environment. One can describe this in terms of stand the nature of neural connections.
In particular, representations of imagined essential nodes27,28. The cortical neural The classification of neuronal inputs is
visions may fail to reach down to the networks (at least for perception) can be still in a primitive state. It is a mistake to
lower echelons of the visual hierarchy. thought of as having nodes. Each node is think of all excitatory neural connections
Coalitions in dreams may be somewhat needed to express one aspect of one per- as the same type. First, connections to a
different from waking ones. cept or another. An aspect cannot become cortical neuron fall roughly into two
10. Penumbra and meaning pathways, but there is little reference to The idea of snapshots is a guess at the
Consider a small set of neurons that fires consciousness. dynamic properties of the parts of a suc-
to, say, some aspect of a face. The experi- There are two fairly recent books cessful coalition, as coalitions are not sta-
menter can discover what visual features expressing ideas that overlap considerably tic, but constantly changing. The
interest such a set of neurons, but how with ours. The first of these is by Edelman penumbra, on the other hand, is all the
does the brain know what that firing rep- and Tononi23. Their dynamic core is neural activity produced by the current
resents? This is the problem of meaning very similar to our coalitions. They also NCC, yet not strictly part of it.
in its broadest sense. divide consciousness into primary con- We also speculate that the actual NCC
2003 Nature Publishing Group http://www.nature.com/natureneuroscience
The NCC at any one time will only sciousness (which is what we are mainly may be expressed by only a small set of
directly involve a fraction of all pyrami- concerned with) and higher-order con- neurons, in particular those that project
dal cells, but this firing will influence sciousness (which we have, for the from the back of cortex to those parts of
many neurons that are not part of the moment, put on one side). They state the front of cortex that are not purely
NCC. These we call the penumbra. The strongly, however, that they dont think motor and that receive feedback from
penumbra consists of both synaptic effects there is a special subset of neurons that there. However, there is much neural
and also firing rates. The penumbra is not alone expresses the NCC. activity leading up to and supporting the
the result of just the sum of the effects of The second book54 is by Bachmann, NCC, so it is important to study this as
each essential node separately, but the who for many years has used the term well as the NCC proper. Moreover, dis-
effects of that NCC as a whole. This microgenesis to mean what is happen- covering the temporal sequence of such
penumbra includes past associations of ing in the 100200 ms leading up to con- activities (A precedes B) will help us to
NCC neurons, the expected consequences sciousness, which is our own main concern move from correlation to causation.
of the NCC, movements (or at least pos- as well. He considers carefully the relevant The explanation here of this inter-
sible plans for movement) associated with psychological phenomena, such as the dif- locking set of ideas is necessarily abbrevi-
NCC neurons, and so on. For example, a ferent types of masking, but has fewer ideas ated. We have outlined some of this in
hammer represented in the NCC is likely about the detailed behavior of neurons. more detail elsewhere10. A more extend-
to influence plans for hammering. A framework somewhat similar to ed account, together with descriptions of
The penumbra, by definition, is not ours has recently been described by the key experimental data, will be pub-
itself conscious, although part of it may Dehaene and Naccache55, though they do lished in a forthcoming book by C.K.56.
become part of the NCC as the NCC not elaborate on the snapshot hypothesis The above framework is a guide to con-
shifts. Some of the penumbra neurons or on the neural basis of essential nodes. structing more detailed hypotheses so that
may project back to parts of the NCC, or they can be tested against already-existing
its support, and thus help to support the General remarks experimental evidence and, above all, to
NCC. The penumbra neurons may be the Almost all of the above ideas have been suggest new experiments. The aim is to
site of unconscious priming48. mentioned previously, either by us or by couch all such explanations in terms of the
others. We have given references to some behavior of identified neurons and in the
Related ideas of these. We believe that the framework dynamics of very large neural assemblies.
In the last 1520 years, there has been an we have proposed knits all these ideas
immense flood of books and papers together, so that for the first time we have Future experiments
about consciousness. For an extensive a coherent scheme for the NCC in philo- These fall under several headings. Much
bibliography, see T. Metzingers home- sophical, psychological and neural terms. further experimental work on small
page: www.philosophie.uni-mainz.de/ What ties all these suggestions togeth- groups of neurons is required for cases in
metzinger. Many people have said that er is the idea of competing coalitions. The which the percept differs significantly from
consciousness is global or has a unity illusion of a homunculus inside the head the sensory input, such as in binocular
(whatever that is), but have provided few looking at the sensory activities of the rivalry57 and in the many visual illusions58.
details about such unity 49 . For many brain suggests that the coalition(s) at the Knowledge of the detailed neu-
years, Baars50 has argued that conscious- back are in some way distinct from the roanatomy of the cerebral cortex needs to
ness must be widely distributed. coalition(s) at the front. The two types of be greatly expanded, in particular to char-
We are not receptive to physicists try- coalitions interact extensively, but not acterize the many different types of pyra-
ing to apply exotic physics to the brain, exactly reciprocally. midal cells in any particular cortical area.
about which they seem to know very lit- Zombie modes show that not all What do they look like, where do they pro-
tle, and even less about consciousness. motor outputs from the cortex are carried ject and, eventually, does each have a set
Dennett 51,52 has written at length out consciously. Consciousness depends of characteristic genetic markers? Are there
about his ideas of multiple drafts, often on certain coalitions that rest on the prop- types of pyramidal cells that do not occur
using elaborate analogies, but he seems not erties of very elaborate neural networks. in all cortical areas? When recording spik-
to believe in the existence of consciousness We consider attention to consist of mech- ing activity from a neuron, it would be
in the same way as we do. Dennett does anisms that bias the competition among very desirable to know what type it is, and
consider a limited number of psychologi- these nascent coalitions. to where this particular cell projects.
cal experiments, but neurons, he tells us, We suggest that each node in these The anatomical methods to character-
are not my department (pers. comm.). networks has a characteristic behavior. ize types of neurons for the macaque
Grossberg53 has written for many years We speculate that the smallest group of monkey have been available for some
about his adaptive resonance theory neurons to be worth considering as a years39 (Fig. 2). In the last two decades,
(ART). This he has developed using very node is a cortical column, with its own very little work has been carried out on
simple neural models. ART involves inter- characteristic behavior (its receptive and cell types and their connectivity, mainly
actions between the forward and the back projective fields). for lack of funds, since such work is not
(for example, to localize seizure onset Health, the National Institute of Mental Health 20. Crick, F. & Koch, C. The unconscious
areas in epileptic patients). With their and the National Science Foundation for homunculus. Neuro-psychoanalysis 2, 311
and subsequent pages for multi-authored
consent, this can provide sparse but criti- financial support. discussion (2000).
cal data about the behavior of neurons 21. Rossetti, Y. Implicit short-lived motor
during conscious perception or imagery59. RECEIVED 12 SEPTEMBER; ACCEPTED representations of space in brain damaged and
It would be very valuable if cortical tissue 9 DECEMBER 2002 healthy subjects. Conscious. Cogn. 7, 520558
(1998).
could be stimulated appropriately with
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