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JOll11lal ofConsciolisness Studies, 1, No.1 , Summer 1994, pp.

91 - 11 8
QUANTUM COHERENCE IN MlCROTUBULES:
A NEURAL BASIS FOR EMERGENT CONSCIOUSNESS?
Stuart R. HamerojJ2
Advanced Biotechnology Laboratory, Department of Anesthesiology, University
of Arizona Health Sciences Center, Tucson, AZ 85724, USA
Abstract: The paper begins with a general introduction to the nature of human consciousness
and outlines several different philosophical approaches. A cr itique of traditional reductioni st
and duali st positi ons is offered and it is suggested that consciousness should be viewed as an
emergent property of physical systems. However, although consciousness has its origin in
di stributed brain processes it has macroscopic properties - most notably the ' unitary sense
of se lf , non-determini stic free will , and non-algorithmic ' intuiti ve' processing - which can
best be described by quantum-mechanical principles.
There have been many attempts by physicists and philosophers to apply quantum-
mechanical principles to the realm of consciousness, but they have been primarily speculative
and analogica l as there has been littl e supportive evidence from the neurosc iences. Thi s paper
postulates that cytoskeleta l microtubules with in neurons could be a possibl e site for quantum
effects.
Although the supportive evidence for such a thesis is, of necessity, highly technical, the
articl e has been structured in such a way that the genera l arguments are accessibl e to the
genera l reader. For those more technically minded, full supportive evidence is avail able in
footnotes and append ices.
Introduction:
Emergence of Consciousness
The nature of human consciousness is the greatest frontier facing science and mankind
and is being approached by many di s ciplines - inc luding neuroscience, cognit ive
science, computer science, phys ics, mathematics, medicine, phannacology, philosophy
and religion. Within thi s di verse approach exists a spectrum which, at one extreme,
cons iders consciousness as s imply the sum total of brain act ivit ies. Thi s reductionist view
is best exemplified by funetionali st philosophers (e.g. Dennett, 199 1) and proponents of
'strong art ifiei al intelligenee' or 'strong AI' (e.g. Minsky, 1986) who eontend that all
mental events may be reduced to an a lgorithm and that a ll brain functions, inc luding
consciousness, may eventua ll y occur in computers . On the other end of the spectrum is a
dualist v iew which cons iders the brain and mind as di stinct , separate entiti es and
eonseiousness as exist ing in a separate reality (e.g. Popper and Eeeles, 1977). Thi s
reduetioni sm/duali sm diehotomy may potentially be resolved by views whieh eontend
that consciousness has a distinct quality, but one which emerges from brain processes
which can be accounted for by natural science.
One aspect of natural science, quantum theory, may be especiall y relevant. In particu-
lar, the concept of quantum coherence leading to macroscopic quantum s tates may help
1 Reprinted in part in the proceedings from the Abisko (Sweden) Conference on Matter Matters -
On the Material Basis of the Cognitive Activity of the Mind, ed. RolfWasen, Peter Arhem, and Uno
Svedin.
2 I am indebted to Alwyn Scott for guidance and suggestions, to Roger Penrose for validation and
inspiration, to Richard Hofstad for patience and clerical talent, and to Gordon Globus, Jane Clark,
Keith Sutherl and and the journal's referees for review, cr iti cism and restructuring. Supported in part
by NSF Grant No. DMS-9 114503. Summer 1994 support from M1DIT, Dani sh Technical Un iversity
is also appreciated.
92 S.R. HAMEROFF
expl ain several of the qualiti es of consciousness which arc difficult to expl ain by
conventional neuroscience. These properti es, and their possibl e quantum correlates (in
parentheses) are:
The ' binding probl em' or ' unitary sense of self' (quantum eoherenee, non-Ioeality).
Randomness, non-detennini sti c free will (quantum indctcnninacy).
Non- algorithmi c ' intuiti ve' processing (computing via quantum superposition).
Reversibl e abl ation by general anaesthesia (prevention of quantum coherence by
anaesthetie inhibition of quantum mobility in protein hydrophobi e regions).
Difference (and transition) between prc-, sub- and non-consc ious processes and
consciousness (collapse of the wave function).
This paper describes a potenti al ' bottom-l eve l' mechani sm in the brain 's recursive,
hi erarehieal organization driven or seleeted by (and also dri ving and seleeting) higher-
level mechani sms (e.g. neural networks, attentional scanning circuits, coherent firing of
di stributed neurons). At thi s ' bottom' level, quantum events can couple directly to
confonnational states of cell proteins. Quantum coherence in cytoskeletal microtubul es
and assoc iated water within each of the brain's neurons may be a bottom level from which
conscIOusness can emerge.
Emergence impli es a qualitatively new property or phenomenon which appears at a
hi erarchical level above the leve l at which rul es of interaction are impl emented. Emergent
properti es (including chaos) occur in many compl ex systems including networks, vor-
tices, lasers, chemical oscillations, genetic networks, weather and deve lopmental pat-
terns. Numerous authors have described consciousness as an emergent phenomenon
ari sing from a hi erarchy of lower-l evel brain processes.
3
Emergent phenomena can exert
downward modulation on lower-l evel interactions to provide reverberation, feedback,
error correction and reward. In the brain's hi erarchy, consciousness (and sub-conscious
faetors) ean regul ate, by downward eontrol , volitional aetions and bodil y funetions.
Consciousness also appears to have emerged at some point in evolution. Perhaps
occurring initi all y as a ' helpl ess-spectator ' epiphenomenon, consciousness then assumed
control of its biological environment (Jaynes, 1976). The emergence of consciousness in
our brains (during each conscious moment, during evolution and during the development
ofeaeh human being) may be likened to new properti es of materials whieh deve lop from
mi croscopi c or quantum-level events. For exampl e, the di stinct properti es of super-
conductivity and superfluidity emerge from materi als as their indi vidual atoms reach a
hi gh level of coherence. In these cases, ordered ali gnment or coherence is due to lowering
temperature to near absolute zero to reduce thermal oscillations. Consequentl y, at a
critical degree of coherence, totally new macroscopi c properti es (superconductivity,
superfluidity) emerge (Leggett , 1989).
Thi s paper suggests that consciousness may emerge as a macroscopic quantum state
from a critical leve l of coherence of quantum-level events in and around a specifi c class
of neurobiological mi crostructure: cytoskeletal microtubules within neurons throughout
the brain.
3 Atkin ( 1992) has defined consciousness as emergent information itself at the moment of its
generati on - ongoing, self-organizing change in a self/ world model. Baas ( 1994) has mathemati-
ca ll y descr ibed ' hyperstructures' with intrinsic observer functions whi ch emerge from hi erarchi es,
and hi erarchi cal schemes of brain organizati on leading to consciousness have been proposed (e.g.
Scott, 1978, 1994; Somj en, 1983) . Globus (1992) descr ibes the brain as ' fractal-like' , with recursive
levels, each using ' edge of chaos' non-linear dynami c processing with ongoing tuning/detuning.
QUANTUM COHERENCE IN MICROTUBULES
Microtubule Associated
Protein (MAP)
Microtubule
Centriole
Membrane
93
Figure 1. Schemati c of simpl e cell with cytoskeletal microtubules organ ized radially around
centr iole near nucleus. Microtubul e associated (MAPs) interconnect mi crotubul es to for m
Dendrit e
Dendritic Spi ne/Synapt ic Receptor
Nucleus
'-.,--/
Membrane L Axon
Microtubule
'-_.n L---- Microtubule Associated Protein
Figure 2. Schematic of centra l region of neuron (d istal axon and dendr ites not shown). Micro-
tubules, interconnected by MAPs, are arranged in parallel. Linking proteins connect mi crotubul es
to membrane prote ins inc luding synaptic receptors on dendritic spines.
Mi crotubul cs (MTs) arc cylindri cal protcin polymcrs intcrconncctcd by crossbridging
protcins (MAPs), whi ch structurally and dynamically organizc functional act i viti cs in
li ving cell s (see Figure I) , including synaptic regulation inside the brain's neurons (see
Figure 2). In Sect ions II and III , we look at some of the current model s of cogniti ve
processes and make a ease for the relevanee of subneuronal struetures. The basie thesis
is that brain-wide microtubule-based quantum states, when coupl ed to synapt ic events
94 S. R. HAMEROFF
and neuronal firing, can help account for the five properti es of consciousness hi ghli ghted
on page 92, and can provide a quantum/c lassical (mindlbrain) interface.
n
Organization of Consciousness within the Brain
The organizati on of consciousness has been approached along three general lines: I)
neural networks, 2) attenti onal scanning circuits, and 3) coherent acti vity of di stributed
neurons.
Neural networks
Neural networks ori ginated from the work of McCull och and Pitts ( 1943) describing
threshold logic neurons and from Hebb 's ( 1949; 1980) concept of the 'cell assembl y' : a
three-dimensional array of interconnected neurons whose firing was predi cted to corre-
late with a spec ific mental state.
4
The neuronal ce ll assembl y concept helped to lead the way to 'artifi cial neural
networks' (' ANNs' ) - parall el, di stributed computer systems designed to impl ement
aspects of brain functi on (e.g. Hopfie ld, 1982). Weighted connecti ons ('synapses')
among processing elements (' neurons') provide ANNs with ' brain-like' learning. pattern
recogniti on and self-organizati on. Churehl attd and Sejnowski ( 1992) have reviewed how
'ANN-like' parall el di stributed processing can operate in the brain, and ANNs are clearl y
more simil ar to brain functi on than are conventi onal algor ithmi c-based AI schemes.
However, the brain! ANN analogy fa lters in at least two maj or areas. First, rather than
being simpl e threshold logic switches as depi cted in ANNs, neuronal synapses are
complex, dynami c connecti ons (' pl asti city ' ). ' In li ving nets everything is continuall y
fluctuating: connecti on we ights, transfer functi ons, parameters, input , and even the very
connecti vity, without leaving any traces of the fluctuati on' (Globus, 1992). Many of these
fl uctuating functi ons depend on the intra-neuronal cytoskeleton; for exampl e Friedrich
( 1990) has shown that learning by increased synapti c efficacy requires eytoskeletal
restructuring in the sub- synapti c region.
It has been claimed that the second maj or di fference between ANNs and brain structure
is that many types of ANNs (e.g. back-propagati on, adapti ve resonance, RCE, sigma pi ,
etc.) use internal process ing within each ' neural element ' . For exampl e in ANNs such as
the useful back-propagati on type, resulting outputs are compared to expected outputs and
the di fference, or error, is conveyed back to an input layer. Werbos ( 1974) ori ginated
ANN back-propagati on algorithms based loosely on Freud 's noti on of retrograde fl ow of
' psychi c energy ' serving feedback/reward functi ons. Because the back-propagati on of
error is counter to unidirecti onal membrane depolari zati on and synapti c transmi ssion,
these types of ANNs have been considered bi ologicall y irrelevant. However retrograde
signalling is now known to occur across synapses (dendrite to axon via ni tric oxide and
other medi ators (e.g. Barinaga, 199 1 , and retrograde infonnati on fl ow within neurons
could be conveyed within mi erotubul es and other eytoskeletal structures (Dayhoff et af.
1994; Rasmussen et af. , 1990). Thus cognitive processes may extend below the level of
synapti c connecti oni sm and include molecul ar-l eve l processing in the cytoskeleton.
4 Hebb estimated that simple assembli es fi re for 1 to 5 milli seconds, more complex ' assembli es of
assembli es' for a half-second, and a ' phase cycle' (a seri es of assembli es of assembli es) for ten
seconds or more (Freeman 1975; Scott 1994). Repeated fir ing of any assembly would lower its
threshold and result in leaming or recognition as the assembly adapts to become more sensitive (by
loweri ng of synaptic threshold - synaptic plasticity) to a given input.
QUANTUM COHERENCE IN MICROTUBULES 95
Attentional scanning circuits and the problem of qualia
Where in the brain docs consciousness occur? Earl y theori es of consciousness invoked a
single brain locati on of self, observation and control, now characteri zed as the search for
the 'grandmother neuron'. Thi s focali zed concept is often generali zed to a 'homunculus'
- the ' litt le man' inside the brain whi ch observes, cont rols and represents the 'self'. But
where in the brain does the homunculus reside? Descartes proposed the pineal gland as a
unitary focus of sclf (although acting as a channel fo r a duali sti c mind entity) partl y
because it was the onl y single, midline brain structure. In hi s functi onali st argument ,
Dennett ( 199 1) criti cizes such a 'Cartesian theater' within the brain whose 'audi ence'
observes and contro ls.
Contrary to these focal models, the evidence gathered from many avenues of research
- stemming from Lashl ey's ( 1950) demonstrati on of the non-l ocali zed di stributi on of
memory - suggests that aspects of consciousness are distributed throughout wide areas
of the brain. Somehow, parall el- arrayed, di stribut ed neuronal groups - perhaps analo-
gous to Hebb's assembli es of assembli es (e.g. ' modul es', 'cartels' etc. (Freeman, 1975;
Gazzani ga, 1985; Minsky, 1986) ) - give ri se to a seri al, integrated stream of conscious-
ness (Baars, 1993). Certain brain and bra instem areas appear to be involved in the
regul ati on, control and focusing of attenti on and consciousness
5
Baars ( 1988; 1993) has
fortnali zed a 'global workspace' model in whi ch multipl e brain ' processors' compete for
access to a broadcasting capability whi ch disseminates the 'winner's' informati on glob-
all y throughout the brain6 Adding the brain stem/ limbi c ' hedoni c' system and language
centres to the external thalamo-corti cal system, Edelman ( 1989) describes re-entrant
circui try from whi ch global mappings of neuronal popul ati ons are 'selected' to reach
consciousness by a Darwini an-like surv ival mechani sm.
Mappings of neuronal acti vity whi ch correlate with a specific mental event still fail to
describe the content, meaning or subj ecti ve sense of self and observer/controll er inherent
in consc iousness. These issues are addressed in Searl e's (1 980) famous 'Chinese Room'
thought experiment (a man sorting Chinese characters) whi ch di stingui shes syntacti c
(l ogical, algorithmi c, comput er-like) from semanti c interpretati on (meaning, under-
standing, human consciousness-like). Taylor ( 1992) observes that content may onl y be
relevant when compared to previous memory: 'the meaning of an input . .. is given by
the degree of overl ap such an input has to past inputs.' But how do past inputs have
meaning? What is the substance of qualia: ' the phenomenal qualiti es of the things of
whi ch we are consc ious, the raw feels and se nsati ons that make up much of our consc ious
li ves' (Korb, 1993).
Functi onali sts like Dennett ( 199 1) argue that quali a are illusory, whil e duali sts beli eve
they are ' ineffabl e' and cannot be understood. Can the redness of a red obj ect be
expl ained by a given pattern of neuronal fi ring, perhaps accented by a neurochemi cal
bathing of a parti cul ar transmitter concocti on? Do two peopl e experi encing red have the
5 These include the reticul ar acti vating system whi ch regul ates wakefulness, the nucl eus reticul aris
of the thalamus whi ch directs attention to di fferent regions of the brain (Scheibel , 1980; Crick,
1984), and the superior colli culus of the tectum whi ch integrates sensory infonnation (Strehler,
199 1). Thalamo-corti cal projections, along with rec iprocal cortico-thalami c feedback have been
impli cated in directing att ention to specific brain regions by 'reentrant feedback loops' (Ede lman,
1989) . Thalamo-corti co-thalami c feedback loops have been observed whi ch sweep from frontal to
occipital cortex approx imately every 12.5 msec. and indi cate brain-wide scanning at a frequency of
about 80 Hz. (Ribary et al., 1991).
6 For example, a cortical focus whi ch by thalamo-cortical feedback is suffic ientl y sel f-sustaining
may be broadcast 'brain-wide' by associative pathways .
96 S.R. HAMEROFF
same sense of redness in their minds? Arc we littl e better than Searle's uncomprehending
man (or Dennett's 'zombi es') with only the illusion of senti ent consciousness? Or do our
senses of beauty, oneness, joy and despair come from another realm? Is there a middl e
ground between reductioni sm and duali sm? Once again quantum coherence at the molec-
ular level of the cytoskeleton offers a poss ibility. Consider the quale of redness; red is
'merely' a specific frequency of the electromagnetic spectrum, a brand of photon. As will
be described, the theory of quantum coherence in the cytoskeleton impli es excitations of
a ground state yielding a condensed fi eld of coherent bosons (phonons, photons, etc.)
with spec ific frequency characteristics. Red of the electromagnetic spectrum can then
correlate with a specific frequency boson-condensed field excitation in the brain. If true,
and when technologically possibl e, the frequency of boson-fi eld exc itations correspond-
ing to red in two people could be compared. More complex qualia (e.g. love, a pepperoni
pi zza, etc.) can then be viewed as complex mixtures of coherent boson fields, which as
macroscopic quantum states, may also have suitabl e properties for a unitary sense of self.
These quantum features , unlike neural firing correlates, convey more than a reduction-
ist representation of sentience; quantum boson fields are unitary, though non-local states.
Their components are indi stingui shabl e from one another by any measuring technique
and they can exist in superposition of many states, collapsing to a singl e one. To which
state a superpositioned quantum wave function collapses can be seemingly random and
'non-detennini st ic '. Thus macroscopic quantum states, unlike neural firing-level corre-
lates, can have non-detennini st ic properti es suggest ive of 'free will ' .
Coherent activity of distributed neurons - the 'binding' problem
Current neurobiological approaches to explain how brain-wide activities result In a
singular perceptual entity involve coherent firing of widely di stributed neurons. Based
on Hebb's cell assembly theory, Milner (1974) proposed that different neurons respond-
ing to a common 'figure' fire syncll1vnously. Using multi-unit recording in awake
monkeys, Abeles (1982) found that temporal correlation among groups of neurons were
related to high-l eve l perceptual tasks and formalized a 'synfire chain' model of cortex.
Von der Marlsburg and Schneider (1986) showed how temporal synchroni zation of
subsets of simultaneously active neurons could 'segment ' different mental objects (e.g.
two different speakers at a noi sy cocktail party) by a postulated fast (ten to hundreds of
milli seconds) Hebbian synaptic modulation mechani sm. (Such rapid synaptic modulation
requires cooperative activities of the intra-neuronal cytoskeleton.) Following Freeman's
(1978) suggesti ve findings in olfactory cortex, Eckhorn ef al. (1988) and Gray and Singer
(1989) found rapid oscillations above EEG frequencies (40 Hz) in cat visual cortex and
Gray ef al. (1989) then showed that these oscillatory responses can become synchroni zed
in a stimulus-dependent manner. Based on these findings, Crick and Koch (1990)
fonnali zed a theory of consciousness in which coherent firing of widely-distributed
neurons in the range of 40 Hz 'bind' them together in mental representation and a unitary
sense of self (see also, Koch, 1993). Studies of auditory-evoked responses in humans
have shown inhibition of 40 Hz coherence with loss of consciousness due to induction of
general anaesthesia (Madler and Poppel , 1987; Plourde and Picton, 1990).
Coherent neural acti vity raises the possibility of a holographic paradigm. Pribram
(1971) proposed that interference of coherent electromagnetic fi elds at the level of
dendritic-dendritic interactions results in a holographic mechani sm of cognitive repre-
sentation. The holographic paradigm is appealing because memory can be recalled by a
'reference' signal , and vast amounts of informat ion can be stored in a di stributed rather
than locali zed manner. Quantum coherence, as proposed to occur in cytoskeletal micro-
QUANTUM COHERENCE IN MICROTUBULES 97
Figure 3.
Rebounding
reaction of
Paramecia
towards external
stimulus. ( 1- 3)
short lasting
cili ary reversal;
(3- 5) pivoting
and circling; (6)
swimming in
new direction.
(With
penni ssion from
Dryl , 1974)
tubules, can provide a mechani sm for intracellular quantum hol ography (Hameroff, 1987;
Schempp, 1993).
Neurobiological approaches (neural networks, attcntional mechani sms and/or coherent
act ivity) generall y describe patterns of neural firing which may correlate with mental
states, including consciousness. However, even if the exact patterns of neuronal firing
correlat ing with specific mental act iviti es become known, would consciousness be better
understood? The reductioni st view would be yes and strong AI advocates would be
validated. To manifest consciousness, computers would merely need to reach the connec-
tioni st complexity of the brain's neurons and synapses - a feat strong AI advocates
believe attai nable within decades. The view of duali sts would be emphat icall y no; the
firing pattern is the brain's act ivity, but it is clearly not equivalent to consciousness.
Emergence partially resolves thi s question; consciousness may indeed be a phenomenon
derived from, but not equi valent to the brain's act ivity. But what exact ly is conscious-
ness? How are di stributed neural act ivit ies, even coherentl y synchroni zed act ivit ies,
bound into a unitary sense of self? Macroscopic quantum states offer one soluti on; but
how could they be supported? Where and how could they exist?
III
Cognition within Living Cells
One poss ibl e path to solving thi s question is by examining the complexity of neurons and
their synapt ic connect ions. Reductioni sts tend to overlook the fact that neurons are alive
and most views of the hi erarchi cal organi zation of the brain stop at the synapse as the
fundamental switch, analogous to states or bits in computers. The complexity of neurons
and their synapses, however, are closer to entire computers than indi vidual switches. Thi s
impli es the mechani sm of consciousness may depend on an understanding of the organi-
zat ion of adapt ive ('cogniti ve') funct ions within li ving cell s.
Like the rest of our ce ll s, neurons are eukaryot ic cell s whi ch, unlike prokaryotic
bacteria, have a true nucleus and cytoskeleton and exhibit mitotic cell divi sion.
7
Single
eukaryot ic cell organisms (protozoa) such as paramecia show remarkable, seemingly
intelli gent abiliti es without the benefit of a single synapse (Figure 3)! Observat ion of
7 Accord ing to Marguli s ( 1975), eukaryotic li fe is the result ofa symbiotic merger ofprokaryotes
( immobil e chemica l factories) and spirochetal invaders (hi ghly motile flagellates with minimal
98 S.R. HAMEROFF
an.t er ior
post eri or
Figure 4a: Scanning
electron mi crograph (rapid
fixation) of Paramecium
showing protruding cili a.
Arrows demonstrate
cili ary movement in
metachronal waves.
4b: Diagram showing
Paramecium polygonal
infrac ili ary lattice with
cili a (ci), containing 9 + 2
sets of mi cro tubules
including peripheral
mi crotubul e (pm)
doubl ets. (ms: medi an
septum, al: alveoli, tt:
trichocyst tips, ps:
para somal sacs, ff: fine
fibri Is, kf: kinetodesmal
fibrils.) This illustrates
how cili a (sensory and
motor) are connected to
intra-cellul ar cytoskeleton.
(With permission from
Wichtennan, 1985).
amoebae hunting food and paramec ia avoiding obstacles led Charles Darwin and other
19th century biologists to project human psychology and aspects of consciousness onto
protozoa. Popul ar books of that era included The Psychic Life of Microorgonisms by
Alfred Binet and The Animal Mind by M.F. Washburn. Darwin viewed adapti ve act iviti es
of singl e-ce ll organisms as rudimentary consciousness related to man through evolution.
Others (e.g. Jaynes, 1976) objected to this linkage, stat ing that protozoan behaviour
'depended entirely on physical chemistry rather than introspecti ve psychology'. How-
ever it now seems introspecti ve psychology itse lf may be an emergent function of
physical chemistry (and quantum mechanic s) at some level. The famous neurosc ienti st
C.S. Sherrington (1951) wrote:
Many forms of motil e single celis lead their own independent li ves. They swim and
crawl, they secure food, they conjugate, they multiply. The observer at once says
'they are ali ve'; the amoeba, paramecium, vorticell a, and so on. They have special-
ized parts for movement, hair-like, whip-like, spiral and spring-like. Sense organs,
beyond a pigment spot, seem to inspection wanting. Of nerve there is no trace. But
the cell framework, the cyto-skeleton. might serve. There is therefore, for such mind
as might be there, no need for our imagination to call halt and say ' the apparatus
for it is wanting'.
As Sherrington observed, the cytoske leton may act as the nervous system of single-ce ll
organisms. Paramecia, for example, can apparently leam (Gelber, 1958), remember
(Applewhite, 1979) and exhibit adapti ve responses such as avoidance and habituat ion
chemi stry) . The motil e spirochetes are thought to have invaded prokaryotes and found shelter and
biochemical energy. The filamentous proteins which compri sed the spirochetal flagellae and pro-
vided movement became the eukaryotic cytoskeleton and offered the new type of cell abiliti es to
1) divide chromosomes and thus mut ate and evolve, 2) compartmentalize, lead ing to specialization,
3) move around, explore and avoid predators, 4) grow asymmetr icall y (e.g. neurons) , and 5)
generall y adapt.
QUANTUM COHERENCE IN MICROTUBULES 99
whi ch involve movement pcrfonncd by coordi nated act ions ('metachronal waves') of
hundreds of hair- like appendages call ed cili a (Figure 4) 8
External events whi ch initi ate adapt ive re sponses via ci li ary and/or membrane pertur-
bation in paramecium are simil ar to receptor binding of neurotransmitter molecules in
neuronal synapses.
9
In paramecia, adapt ive behavioural responses to external st imuli
involve motor act iviti es performed by complex act iviti es of cili a; in neurons they include
regulation of synapt ic strengths. and cogniti ve funct ions related to learning. memory, and
aspects of consc iousness.
Is consciousness a property of protoplasm in general? Do single-cell organisms, such
as paramecium, exhibit consciousness? If not, at what point in the evoluti onary hi erarchy
does consciousness emerge? These di stinctions may be somewhat arbitrary, but the/abric
of consciousness may be present within a ll eukaryot ic ce ll s. If so, how low in the
organi zat ional hi erarchy of single cell s do the rudiments of consciousness extend? An
extreme view ('panpsychi sm') is that consciousness is a quality of all matter: atoms and
their subatomic components having elements of consciousness (Spinoza, 1677; Rensch,
1960). But then what di stingui shes non-li ving material from li vi ng, and how are all the
atomic and sub-atomic consciousness entiti es unifi ed (the binding problem taken to
extreme)?
Many contemporary thinkers fee l that quantum mechanics may provide answers to at
least some of these questions and that consc iousness may emerge at a crit ical amount of
non-l ocal quantum processing linked to some neural structure. It is the particular proposal
of thi s paper, deve loped in the next section, that cytoskeletal microtubul es are the most
likely locat ion for such an effect. Preliminary technical arguments are developed in
Appendices 1-3. Appendix 1 di scusses the structure of micro tubules in detail and reviews
recent evidence for their role in cogniti on; Appendix 2 out lines Frohli ch's work on
protein dynamics, whil st Appendix 3 outlines the mechani sms by whi ch proteins may act
as complex infonnat ion processors and 'quantum computers ' . The arguments are, of
necessity, somewhat technical. Readers with a scientifi c background should read the
appendi ces in full at thi s stage, whil e others may prefer to proceed to the next section.
8 Compr ised of nine microtubule doublets arranged in a ring around a central microtubule pa ir, cili a
are membrane-bound extensions of the cytoskeleton in cell s ranging from protozoa to human
epithelium. In additi on to several types of motor movement, cili a have sensory function in that their
perturbat ion is transmitted to the cell. Lund ( 1933 ) showed that microtubules withi n paramecia were
' conductil e' and transmitted information, and Atema ( 1974) proposed that signal transduction in
sensory cili a involved propagating confonnationa l changes along cili ary microtubule subunits.
Complex action of paramecium cili ary ' metachronal waves' are coord inated by a per ipheral cyto-
skeletal network to wh ich the cili a are anchored; thi s network is in tum connected to intemal
microtubules and other cytoskeletal structures. Several authors (e.g. Lund, 1933; 194 1) identified a
centra l confl uence of cytoskeletal structures (the ' neuromotorium' ) characterized as the paramecium
' brain ' . Metachrona l waves of cili ary beating in paramecia are reversibly inhi bited by the general
anaesthetic chl orofonn (parducz, 1962), suggesting some link to consc iousness in hi gher organ isms.
9 Each triggers 'second messenger' transduction cascades (G proteins, calcium ion flux, adenyl
cyclase, phosphatidyli nos itol, cycl ic AMP, etc.) which in turn activate enzymes (e.g. dynein
AT- Pases, cytoplasmic phosphatases and kinases such as calcium-calmodulin prote in kinase, MAP
kinase, calc ineuri n, protein kinases A and C (Er iksson et aI., 1992; Montoro et aI., 1993)) . Among
the responses of the enzymes (kinases in particul ar) are phosphorylation/dephosphorylation of
cytoskeletal structures including microtubule-associated-prote ins ('MAPs' ) and intenned iate
(neuro) filament s wh ich can initi ate cytoske letal s igna lling and structural reconfiguration (Johnson
and Pope, 1992). In additi on, external events acting through membrane prote ins can directly connect
to the interna l cytoskeleton by linking prote ins such as fodrin, ankyrin and actin.
100 S.R. HAMEROFF
IV
Quantum Physics and Consciousness
Quantum theory describes fundamental properti es ofmattcr and energy which comprise
our uni verse. Seemingly bi zarre and counter-intuitive, quantum properti es - including
wave/particl e duality, the (Heisenberg) ' uncertainty principl e', randomness or acausality,
and quantum inseparability or non-Ioeality (quantum eoherenee) - have been repeatedl y
verifi ed by experimentation over much of thi s century. These strange properti es may help
expl ain the mysterious nature of consciousness. ' Perhaps our minds are qualiti es rooted
in some strange and wonderful feature of those physieal laws whieh aetuall y govern the
world we inhabit ' (Penrose, 1989).
At the base of quantum theory is the wave/partiel e duality of atoms and their eompo-
ncnts. When unobserved, an atom or sub-atomic particl e behaves as a 'wave ofpossibili-
ti es '; observation in effect 'coll apses the wave function ' and a particl e appears. The
Heisenberg uncertainty principl e dictates that either the position or momentum of a
partiel e (but not both) ean be known with arbitrary preeision. Beginning over four
decades ago, these properti es prompted very general comparisons to mental processes.
As observation of a quantum partiel e alters its state, Bohm (1952) observed that thoughts
and mental processes, when focused upon, become altered and might therefore have some
basis in quantum meehanies. Szent-Gyorgyi (1960) and Pullman and Pullman (1963)
made broad analogies between wave/particl e and mindlbrain dualiti es.
Quantum randomness stems from the superposition of an unobserved atom or particl e
in many possibl e states; which particul ar state will be observed is apparentl y random.
Einstein objected to thi s quantum randomness or acausality: ' I cannot beli eve that God
would pl ay dice with the Universe'. However as Page ls pointed out ' if things at base are
utterl y random, nothing can make them more di sordered. Compl ete randomness at the
heart of things is the most stabl e situation imaginabl e' (Herbert , 1993). Noting that the
' materi al' world consists of nothing but 'relentl essly unreali zed vibratory possibiliti es' ,
von Neumann proposed and Wigner and others elaborated (Wheeler and Zurek, 1983)
that by observing (measuring) the real world, the eonseious mind eollapses wave fune-
tions and eonfers solidity, singl e-valueness and dependabl e eonstaney. Globus (1987)
argues that
possibl e worlds are superposed within the brain - the 'holoworld', and perception,
both waking and dreaming, se leets a world for unfolding from the holoworld. The
coll apse of the wave function is a se lection process from a priori impli cate worlds.
These ideas are eomparable to Bohm's (1980) ' implieate order' , and Everett's (1957)
many uni verse view. But then what is it about the conscious mind which causes thi s
eollapse?
Another view is that matter consists of fl eeting vibratory patterns in a vast fi eld of
consciousness: a uni versal mentality interpenetrates the physical world. This concept
(' quantum animi sm') suggests that every quantum wave contains consciousness and vice
versa: 'where there's a will , there 's a wave' (Herbert, 1993). Thi s view is a quantum
version of duali st and ideali st concepts such as Cartesian mental substance or Hege lian
world spirit (Searle, 1980); the quantum/mental realm interaets with the physieal realm
in the structures of our brain. Several approaches to elucidate the site and mechani sm of
thi s quantum interaction have focused on the synapse.
Beek and Eeel es (1992) proposed that quantum randomness results in what appears to
be probabili stic, acausal synaptic function. Noting that in an apparentl y random fashion
QUANTUM COHERENCE IN MICROTUBULES 101
onl y onc of approximately six axonal acti on potenti als results in synapti c release of
neurotransmitter vesicles, they pinpoint the quantum effect as the ' movement of a
hydrogen bridge by electroni c rearrangement ' acting on vesicle release through the
pre-synapti c vesicul ar grid. A hexagonal latti ce on the inner surface of the presynapti c
region, the vesicul ar grid was chosen by Beck and Ecc les as the site for quantum
influence because of it s ' paracrystalline structure' whi ch ' makes it possibl e to have
long-range interacti ons between the consti tuents', (A simil ar argument can be made for
more ubiquitous paracrystalline mi crotubul e arrays. ) Other candidates for medi ati on of
quantum randomness effeets inelude ion ehannels (Bass, 1975; Donald, 1990) and ions
(e.g. ealeium ions) themse lves (Stapp, 1993).
Quantum coherence
The greatest surpri se to emerge from quantum theory is quantum inseparability or
non-l ocality whi ch impli es that all obj ects that have once interacted are in some sense
still connected! Schrodingcr observed in 1935 that when two quantum systems interact,
their wave functi ons become ' phase entangled'. Consequentl y, when onc system's wave
functi on is coll apsed, the other system's wave functi on, no matt er how far away, instantl y
eoll apses too ( Herbert , 1993). The non-I oeal eonneeti on (quantum eoherenee) is not onl y
instantaneous and independent of di stance but impervious to shi elding. Einstein, Podol-
sky and Rosen ( 1935) proposed an experimental method for testing inseparability involv-
ing observati on of polari zati on states of two photons emitted in di fferent directi ons from
a eommon souree. 1.S. Bell in 1964 (e.g. Bell , 1987) formul ated the noti on that, if
quantum inseparability were fa lse, then changing the measurement context of one photon
would not affeet the other photon's behaviour ('Bell 's theorem' ). Amazingly, experi-
ments by Clauser et af . ( 1969) and Aspeet and Grangier ( 1986) di sproved Bell 's theorem
by showing that choice of context of measurement of one photon instantl y affects
outcome of measurement of a di stant photon. The quantum world is trul y interconnected;
quantum entities are 'aware' of the states of their spatially separated relatives !
Such ' knowing' has prompted linkage of non-l ocal quantum coherence to mental
proeesses. Bohm ( 195 1) deseribed the quantum world as an ' undi vided wholeness' or
' impli cate order ' and related it to a holographi c model of consciousness whi ch accents
the wave nature of experi ence. Pribram ( 197 1) has proposed that mental representati on
oeeurs holographi eall y by interferenee of eo he rent waves (e.g. dendriti e fi eld potenti als).
Non-l ocal quantum effects among di fferent indi viduals have been related to ' Jungian
synehroni eity' (, meaningful eoineidenee') ori ginating from what the famous psyeho-
analyst e.G. lung termed the 'eoll eeti ve uneonse ious' ( Peat, 1992; Insinna, 1992).
Quantum coherence has also been linked to more spec ifi c bi ological functi on and
eomputati on. Conrad ( 1992) has argued that superpositi on of spati all y-separated eIeetron
states fac ilitates common bi omolecul ar int eracti ons such as anti gen-antibody, neuro-
transmitter-receptor, enzyme- substrate, protein se lf-assembl y and other functi ons. Con-
rad further suggests that quantum coherence and parall eli sm inherent in the quantum-
mechani cal wave functi on can enhance the speed, reli ability, and energy di ssipati on of
elementary switches such as those involved in molecul ar computati on (i ncluding, per-
haps, tubulins within mi crotubul es). Theoreti cal descripti ons of quantum computers
propose utili zing quantum coherence, superpositi on and parall eli sm for technological
purposes.'o Penrose ( 1989; 1994) sees sueh quantum eomputing in the brain as a meehan-
10 For example Deut sch ( 1985; 1992) and Ll oyd ( 1993) forsee quantum-level bit states (e.g. 1,0)
whi ch can be in quantum superpositi on of both I and O. If the bit state sites (in an array or polymer)
102 S.R. HAMEROFF
ism for ' non-algor ithmi c' processing intrinsic to human consciousness. He speculates
that conscious thinking involves resolution of alternat ives that were previously in linear
superposit ion.
Coherent quantum phenomena cannot perform computat ion if the system undergoes
signifi cant ' measurement-like interact ions' with the outside wor ld; onl y well-i solated
quantum computers arc possible. Inherent isolat ion impli es a 'subject ivity', or internal
informat ion inaccessibl e to external observat ion within quantum computers, a property
suggest ive of consciousness (Deutsch, 1992). An appropriate 'classical' interface for
quantum computers could be 'select ive driving of resonances ' (Haddon and St illinger,
1982). Ll oyd ( 1993) proposcs an array ofwcakly-coupl cd quantum systcms whi ch would
be subj ectcd to sequences of clectromagnet ic pul ses of wcll-defi ncd frequency and
Icngth, pcrmitting loadi ng, proccssing and unloading of informat ion. Cytoskclctal poly-
mers subj ected to neuronal firing might constitute such a system.
Penrose ( 1989) has also invoked the poss ibility of quantum superpositi on in synapt ic
plasticity. Non-periodic tiling patterns of quasi- crystals ('Penrose til es' ) require ' non-
local' effects; it appears necessary to 'know' the state of the pattern many til es away from
the point of assembly. Rather than classical crystal growth in whi ch indi vidual units
attach themselves at a continuall y moving growth line, Penrose considers thi s to occur as
an 'evolving quantum linear superpositi on of many different alternat ive arrangements'
and likens thi s situat ion to learning and memory by synapt ic plasticity in whi ch neuronal
connect ions are rapidl y fonned, act ivated or deacti vated. ' Thus not just one of the
possibl e alternat ive arrangements is tried out , but vast numbers, all superposed in
complex linear superpositi on' (Penrose, 1989). Synapt ic conncct ions are formcd and
rcgulatcd by cytoskc lctal polymcrs, including microtubulcs. Complcx lincar supcr-
positi on of many cytoskeletal confi gurat ions corresponding to synapt ic connect ion pat-
terns coll apsing to a single one could account for the extremely rapid synapt ic plasticity
rcquircd for cogniti on. Pcnrosc ( 1989; 1994) also considcrs that coll apsc of supcr-
positi oned states and conversion from quantum to classical behaviour depends on a
threshold related to quantum gravity. He prcdicts that gravity, although an extremely
weak force, influences the quantum realm because it acts on the spacetime structure itse lf.
Objects that become larger than a cr ucial size spontaneously actuali ze one of their
possibiliti es; spacetime curvature causes the system' s wave function to coll apse ' under
its own wc ight ' . Hcrbcrt ( 1993) has uscd a ' quantum rcalm' approach to cst imatc a mass
value for 'quantumness' and obtains a rough estimate of 10
6
daltons. Thi s is the size of
Eccles' pre-synapt ic 'microsite' and also of about ten tubulin subunits within a micro-
tubule. Penrose calcul ates a hi gher number oftubulins whose coherence for a criti cal time
will self-coll apse, and result in a single conscious event. According to Penrose, the
manner in whi ch quantum gravity can act as a bridge between the quantum world and the
classical world wi ll await new di scoveri es in quantum physics. I I
Bose-Einstein condensates
Concluding that quantum effects are necessary to explain the ' unity of conscious experi-
ence' , Marshall ( 1989) has searchcd for a quantum mechani sm that is extended in space
are locall y interactive as in a cellul ar automaton (e.g. from overl ap among uni t- unit electron wave
functions, van der Waals force s, confonnational changes of subunits, dipole coupli ng, etc.), then
parallel logic and quantum coherence can result in quantum computation; multiple computations
coll apse into a macroscopic quantum state.
II See also (in thi s volume) Nunn et.al. ( 1994)
QUANTUM COHERENCE IN MICROTUBULES 103
(non-local , long-range ordcr), capable of many states, and has a 'sharcd idcntity' un-
analyzabl e into parts (,diversity in unity'). Marshall concludes these properti cs are
uniquc to systcms known as 'Bosc-Einstcin condcnsatcs ' (scc Appcndix 2). Usually, each
particl e/wave in a quantum system possesses its own wave function. However, under
certain circumstances, many quantum parti cle/waves move coherently, described by the
same wave function. Arising in quantum fi eld theory, Bose-Einstein condensates are
collective, macroscopic quantum states named after Indian physici st Satyandra Nath
Bose and Albert Einstein who independentl y predicted the types of particl e/waves
(' bosons' ) and conditions necessary for reali zation of such states (Zohar and Marshall ,
1994).
Quantum fi eld theory describes the underlying reality of everything in the uni verse
(including consciousness) as consisting of three components: the vacuum, space and time.
A ' fi eld of fi elds' which contains no particl es. the vacuum gives ri se to quantum
wave/ particl es as exc itations or energy fluctuations within it (like 'sound from a drum
skin' (Zohar, 1990)). A ground state is a component fi eld of the vacuum whose excitation
can yield the two types of wave/particl es: fermions and bosons. Fennions are the
ingredi ents of matter (electrons, protons, and neutrons) which tend to be indi vidual; their
wave functions cannot overlap entirely (although fennion pair wave functions have
bosonic properti es and can overlap as in superconductors, supermagnets and superfluids).
Bosons are wave/particl es of rel ationship which convey forces and which can merge as
their wave functions overlap; they can coherently share identiti es. Bosons include pho-
tons, virtual photons, phonons and other entiti es including possibly gravitons; the forces
they convey include electromagnetic, weak and strong nucl ear and possibly gravity. In
concert with the uncertainty principl e, a set of boson ground-state excitations can act
coherently by entry into the lowest momentum state so that position becomes undefined
(Tilley and Till ey, 1986). Consequently, a large set of bosons is described by a singl e
wave function and coherent , collective macroscopic quantum states result (Herbert ,
1993).
Quantum fi eld theory was initi all y linked to consciousness by Ricci ardi and Umezawa
(1967) who proposed that each brain cell contains a spati all y di stributed system with a
full complement of quantum-mechanical degrees of freedom. Without specifying the
intraneuronal structural correl ate, they suggested that memory of an external stimulus is
imprinted as a perturbation of the vacuum ground state of the unspec ifi ed intraneuronal
system. By viol ating the system' s original dynamical symmetry, long-range correlation
waves (bosons) condense and correlate with mental processes.
Examples of known Bose-Einstein condensates include the laser, in which many
photons occupy exactly the same optical state, superconductors, in which numerous
linked electrons ('Cooper pairs' ) take on identical quantum possibiliti es, supennagnets
in which micro-dipol es ali gn, and superfluids like helium, in which quantum synchro-
ni zed atoms create a friction-free fluid. In each case the overlapping of position gives a
shared identity and the ground state has a long-range order (coherence).
Superconductors, supennagnets and superfluids require temperatures near absolute
zero to reduce thennal oscillations and bring the particl es into coherence. Lasers require
high energy to dri ve crystals into coherent excited states. Of known systems akin to
Bose-Einstein condensates , only Frohlich's proposed pumped phonon system (see
Appendix 2) can operate at body temperatur e (Marshall , 1989). Frohlich predicted that a
set of proteins in a common electromagnetic fi eld (e.g. proteins within a pol ari zed
membrane, subunits within an electret polymer like microtubul es) undergo coherent
conformational excitations if biochemical energy were suppli ed ('pumped phonons' ).
104 S.R. HAMEROFF
Figure 5 Microtubul e automata (MTA) with ordered water.
Using a set of assumpt ions for a layered sheet of dipoles, Clarke ( 1994) has questioned
the feasibility of Frohli ch's biological Bose-Einstein condensates. However cooperat ive
effects of coupl ed, ordered water and non-planar geometries (i n particular cylindri cal
protein latt ices like microtubules) make Frohli ch's biological Bose-Einstein condensates
more likely (e.g. Satarie et al., 1993; Jibu et al., 1994).
Cit ing Frohli ch's pumped phonons in unspecified ' neural proteins' as a Bose-Einstein
eondensate, Marshall predieted that eonseious phenomena (thoughts, images, ete.) would
occur as excitat ions of the ground state, like waves in superfl uid helium or holograms in
laser li ght. He reasoned that such a condensate could suffi ce even as only a minor facet
of brain dynamics (i n a superconductor onl y onc material part in tcn thousand participates
in the phase ehange, but the maeroseopie properties are dramatieally alt ered). Cyto-
skeletal microtubul es have suitable qualifications to act as a brain substrate for Bose-
Ei nstein condensates related to consciousness .12 Consciousness may then be seen as
a dynamic macroscopic quantum state originati ng from a mixture of coherent states
originati ng in microtubules.
Super-radiance
Quantum eoherenee has been theoretieally appli ed not only to membranes and eytoskele-
tal str uctures, but to water molecules ordered at their surfaces (Figure 5).13 Using
12 Froh li ch pumped phonons have already been considered in microtubules as a basis for infonna-
tion processing via spin-glass behaviour and as a clocking mechani sm for cellul ar automata
functions. The cooperative coupling resulti ng in coherence has been assumed to be phonons
(bosons) coupl ed to dipoles or electrons ' trapped' in hydrophobic regions (e.g. aromatic ami no acid
residues with resonant orbitals) . Microtubul es' paracrystalline periodic lattice and cylindri cal
configuration provide an ideal coherent structure, and the functions of microtubules are crucial to
intracellul ar cogn iti on. Thus spin-glass and automata models, simil ar if not equi valent to Bose-
Einstein condensates, may manifest quantum computing by interactive patterns of coherent sets of
pumped phonons (bosons) .
13 Frohli ch theorized that energy could be stored without thermal loss in coherent, dipolar propa-
gating proteins or in a thin layer of water and ions just beneath the cel l membrane. Watterson ( 1987)
has shown how organized, dynamic water clusters ('pixe ls') can represent infonnat ion at membrane
and cytoskeletal surfaces. Consider ing the layer of ordered water out side and inside microtubules,
Del Giudi ce et al. ( 1983) proposed that microtubules' cylindri ca l structure may be understood by
quantum-vacuum symmetry breaking and boson se lf-focusing by ordered water. Like the Meissner
QUANTUM COHERENCE IN MICROTUBULES 105
quantum fi eld theory, Jibu et al. ( 1994) have proposed that ordering of water moleeul es
and the quanti zed electromagneti c fi eld confined inside the holl ow mi crotubul e core
manifest a specifi c coll ecti ve dynami cs call ed super-radiance. Accordingly. each mi cro-
tubul e can transfonn incoherent, di sordered energy (molecul ar, thermal, or electromag-
neti c) into coherent photons within its holl ow core, Furthennore, coherent photons
created by super-radi ance may penetrate without loss along the mi crotubul e as if the
opti cal medium were made ' transparent ' by the propagating photons themselves. Thi s is
a quantum phenomenon eall ed self-induced transparency (MeCull and Hahn, 1987)
(Figure 6). Some evidenee suggests weak photoemi ssion fr om li ving ee ll s, and Popp
( 1986) has proposed a regul atory rol e for 'eoherent bi ophotons'. Thus Jibu et al. suggest
that mi crotubul es can behave as opti cal waveguides whi ch result in coherent photons;
they estimate that thi s quantum coherence is capabl e of superpositi on of states among
mi crotubul es spati all y di stributed over hundreds of mi crons. These in turn are in super-
positi on with other mi crotubul es hundreds of mi crons away in other directi ons and so on.
Consequentl y mi crotubul e quantum dynami cs may be coupl ed over brain- wide areas, a
superpositi on whi eh eould solve the ' binding probl em' and aeeount for unity of thought
and consciousness.
To summari ze, cytoskeletal mi crotubul es are likely candidates for quantum coherence
relevant to consciousness because:
Microtubul e indi vidual subunit (tubulin) confonnati on may be coupl ed to quan-
tum-l evel events (el ectron movement , dipole, phonon) in hydrophobi e protein
regIOns.
Microtubul e paracrystalline latti ce structure, symmetry, cylindri cal configurati on
and parall el ali gnment promote long-range cooperati vity and order.
Holl ow mi crotubul e interi ors appear capabl e of water- ordering, waveguide super-
radiance and self-induced transparency.
It is proposed, therefore, that mi crotubul es' ubiquitous presence and crucial involvement
in regul atory and cogniti ve functi ons in neurons and other eukaryoti c cell s can medi ate
quantum effects in a wide range of signifi cant bi ological functi ons. In parti cul ar, quantum
coherence (Frohli ch pumped phonons, Bose-Einstein condensati on, ordering of water) in
cytoskeletal mi crotubul es and related structures can lead to quantum computing and
emergence of macroscopi c quantum states suitabl e for a unitary sense of consciousness.
V
Anaesthesia, Quantum Coherence and Consciousness
As a putati ve mechani sm for consciousness, mi crotubul e-based quantum coherence
should be sensiti ve to general anaesthesia. Brains of pati ents under general anaesthesia
are commonl y quite acti ve: EEG, evoked potenti als, respiratory and autonomi c dri ves
and other brain functi ons persist despite a lack of consciousness. Thus, at just the ri ght
level, general anaesthesia is the absence of consciousness. 14
effect for superconducti ng medi a, electromagneti c energy would be confi ned ins ide fil amentous
regions around whi ch the mi crotubul e subunits gather. Del Giudi ce's group (1 983) showed that thi s
sel f-foc using should result in fil amentous beams of radius 15 nanometers, precisel y the inner
di ameter of mi cro tubul es!
14 A vari ety of di fferent types of gas molecul es constitute general anaestheti cs; they range from
halogenated hydrocarbons to ethers to inert gases such as xenon. What they share in common are I)
106 S.R. HAMEROFF
If quantum coherence in microtubulcs is essenti al for consciousness, anaesthet ics must
somehow inhibit or prevent it, either indirectly or directly. IS At concentrations of anaes-
thesia just sufficient for ablat ion of consciousness, anaesthetic act ion at hydro- phobic
regions of membrane receptors and channels, tubulins and other proteins could have
profound effects by retarding mobility of dipoles/electrons in those hydrophobic regions,
thereby inhibiting confonnational dynamics. coherent phonons and Bose-Einstein con-
densation. Wulfand Featherstone (1957) showed that anaesthetic-protein binding within
hydrophobic regions alters protein-water binding at the protein surface. Thus anaes-
thetic-tubulin interact ion would alter cooperative water-binding and ordering. with
consequent effects on quantum vacuum symmetry breaking and attendant 'super-
radiance' and 'self-induced transparency'. By having such effects in hydrophobic regions
oftubulin and other proteins, anaesthetics can reduce and prevent quantum superposition
and coherence. This is consistent with the quantum realm interact ing with the brain via a
collection of quantum-level events in hydrophobic regions of brain proteins - in
particular tubulin subunits within microtubules.
VI
Summary
Consciousness is described as an emergent macroscopic quantum state driven or selected
by neurobiological mechanisms (neural networks, attentional scanning circuits, coherent
firing of distributed neurons) with origins in quantum coherence in cytoskeletal micro-
tubules within the brain's neurons. Microtubule quantum coherence is thought to derive
from two possibly inter-related mechanisms:
Bose-Einstein condensates stemming from a 'Frohlich pumped phonon' mecha-
ni sm of dipoles in hydrophobic pockets of microtubule subunits .
A quantum-dynamical system of water molecules and the quantized electromag-
net ic field confined inside the hollow microtubule core. The latter induces a
specific collective dynamics called super-radiance by which coherent photons are
created and penetrate without dissipation in the microtubule core in a quantum
effect called self-induced transparency (Jibu et al. , 1994).
A macroscopic quantum state approaching brain-wide dimension can provide proper-
ties which define consciousness. These include: I) a unitary sense of self, 2) randomness
or non-detennini st ic 'free will ', 3) non- algorithmic 'quantum' computing. Coupling of
solubility in a specifi c ' hydrophobic' environment, and 2) binding by weak van der Waals forces
(e.g. Halsey, 1983). After many years of investigating effects in hydrophobic lipid regions of
membranes, anaesthetic gas mol ecul es are now recognized to reversibly inhibit consciousness by
weak, van der Waals binding in hydrophobic regions of some set of neural (presumably synaptic
membrane) proteins (Franks and Lieb, 1982). As previously described, quantum-level events
(dipole/electron oscill ati ons, phonons) in protein hydrophobic regions are thought to coupl e to
functional dynamic protein confonnational states. Anaesthetics are known to retard mobility of
electrons (Hameroff and Watt, 1983), and by doing so in hydrophobic regions anaesthetics may
prevent protein confonnational responsiveness.
15 Possible indirect ways include inhibition of activiti es of membrane, second messenger or
membrane-cytoskeletal linking proteins wh ich could disconnect and isolate cytoske letal dynamics
from the external mili eu. Anaesthetics may also disrupt hydrophobic links among mi crotubul e-
assoc iated-proteins (MAPs) wh ich interconnect mi crotubul es into functional networks (Lewis et aI. ,
1989). Alternative ly, anaesthetics may directly alter cytoskeletal quantum coherence. General
anaesthetics are known to bind to mi crotubules and at hi gh enough concentrations cause their
depolymerization (Allison and Nunn, 1968) .
QUANTUM COHERENCE IN MICROTUBULES 107
(a)
(b)
(c)
(d)
Figure 6. A schematic representation afthe process afsuper-radiance in a microtubule
Each oval without an arrow stands for a water molecule in the lowest rotational energy state. Each
oval with an arrow stands for a water molecule in the fir st excited rotational energy state. The process
is cycl ic (a), (b), (c), (d), (a), (b) , and so on.
(a) lnitial state of the system of water molecules in a mi crotubul e: energy gain due to the thennal
fluctuation oftubulins increases the number of water molecul es in the fir st excited rotational energy
state.
(b) A coll ective mode of the system of water molecules in rotationally exc ited states: a long-range
coherence is achi eved inside a mi crotubul e by means of spontaneous symmetry breaking.
(c) A coll ective mode of the system of water molecul es in rotationally excit ed states loses its
energy coll ectively, and creates coherent photons in the quantized electromagnetic field inside a
mi crotubul e.
(d) Water molecules, having lost their first excit ed rotational energies by super.rad iance, start
again to gain energy from the thermal fluctuation of tubulins, and the system of water molecul es
recovers the initi al state (a) . (With penni ssion from Jibu et aI. , 1994) .
108 S.R. HAMEROFF
thi s quantum state to mi crotubul e subunit conformati onal states can account for cellul ar
control, adapti ve behaviour, synapti c regul ati on and cogniti on by propagating s ignals
through cytoskeletal networks and infonnati on processing via mechani sms including
moleeul ar (,ee llul ar ' ) automata, spin-glass and holography. Mierotubul es are appropri ate
candidates for quantum coherence leading to consciousness because their subunit confor-
mati on (and consequent functi on) can coupl e to quantum events in hydrophobi c regions.
their paracrystalline latti ce structure and symmetry promote long-range order and their
holl ow cylindri cal core can lead to water ordering with waveguide super-radiance and
self-induced transparency. The essenti al and ubiquitous roles mi crotubul es pl ay in cellu-
lar control and regul ati on and their communi cati on and proximity with membrane
proteins, cell nucleus and other bi omolecul ar structures can link mi crotubul e-based
quantum events to basic bi ological functi ons.
General anaestheti c gas molecul es, whi ch selecti vely inhibit consciousness at appro-
pri ate concentrati ons, act at hydrophobi c protein regions and are known to inhibit
electron mobility. Thus anaestheti cs, whether acting directl y on mi crotubul es or in-
directl y via membrane proteins or membrane- cytoskeletal connecti ons, can reversibl y
erase consciousness by inhibiting quantum-l eve l events in protein hydrophobi c regions
and thus di srupting quantum coherence.
As a model of consciousness, quantum coherence in mi crotubul es is reducti oni st in that
a specifi c molecul ar structure is featured as a site for consciousness. It is seemingly
duali st in that the quantum realm (whi ch is actuall y intrinsic to all of nature) is seen to
act through mi crotubul es. General anaesthesia is viewed as reducing and preventing
quantum superpositi on and coherence in protein hydrophobi c regions, a view whi ch also
speaks to an intrinsic quantum nature of consciousness.
APPENDIX
I
Microtubules and the Cytoskeleton
Interi ors of li ving eukaryoti e eell s are strueturall y and dynami eall y organi zed by net-
works of interconnected protein polymers. Ori ginall y tenned the cytoskeleton because of
their ' bone-like' structural support , these networks also orchestrate and control dynami c
cellul ar acti viti es and may be considered as the cell 's nervous system. The cytoskeleton
consists of mi crotubul es (,MTs' ), actin mi crofi laments, intennedi ate fil aments and an
organi zing compl ex call ed the centrosome whose main components are MT super-
eylinders eall ed eentri oles (Dustin, 1984). Parall el-arrayed MTs are intereonneeted by
eross-bridging proteins (MT-assoe iated proteins: 'MAPs' ) to other MTs, organell es,
fil aments and membranes to fonn dynami c networks. Contractil e MAPs such as dynein
and kinesin parti cipate in cell movement as well as intra-neuronal ('axopl asmi c' ) trans-
port whi ch moves materi al and pl ays a maj or rol e in maintaining and regul ating synapses.
MAPs whi eh form struetural bridges stabilize MTs, prevent their di sassembl y and may
be phosphorylated and impart energy into the eytoskeleton. Thus MAP- MT eytoskeletal
networks determine ce ll architecture and dynami c functi ons (e.g. mitosis, growth, di ffe r-
enti ati on, movement, synapse fonnati on and functi on, etc. ) essenti al to the li ving state.
Of the fil amentous structures whi ch compri se the cytoskeleton, MTs are the best
characteri zed. MTs are holl ow cylinders 25 nanometers (' nm' : I 0-
9
meters) in di ameter
whose wall s are 13 ehains of subunit proteins known as tubulin. Eaeh tubulin subunit is
a polar, 8 nm dimer whi ch consists of two s li ghtl y di ffe rent classes of 4 nm monomers
QUANTUM COHERENCE IN MICROTUBULES 109
nm
Figure 7.
Microtubul e (MT) structure from
X.ray crystall ography (Amos and
Klug, 1974). Tubulin subunits are
8 nm dimers comprised of a and ~
monomers.
(mol ecul ar weight 55,000 daltons) known as a and ~ tubulin (Figure 7). Each dimer (as
we ll as whol e MTs) has a dipol e with negati ve charges locali zed towards a monomers
(DeBrabander, 1982). Thus MTs are 'elec trets': ori ented assembli es of dipol es with
pi ezoelectri c properti es (Athenstaedt , 1974; Masearenhas, 1974). Dimers self-assembl e
into MTs, apparentl y in an entropy-dri ven process whi ch can qui ckl y change by MT
di sassembl y and reassembl y in another ori entation (' dynami c instability' (Kirschner and
Mitehi son, 1986) ). The tubulin dimer subunit s within MTs are arranged in a sli ghtl y
twisted hexagonal latti ce, resulting in differing neighbour relati onships among each sub-
unit and it s six nearest neighbours. Tubulin undergoes confonnati onal changes induced
by hydrolysis of bound GTP to GDP (hi gh energy phosphate molecul es analogous to ATP
and ADP (Enge lborghs 1992; Melki et at., 1989 and bi ochemi cal energy can also be
pumped int o MTs by phosphorylati on/dephosphorylati on of MAPs (Theurkauf and
Vall ee, 1983). Genes for a and ~ tubulin arc compl ex and give ri se to varying tubulin
primary structure. For exampl e. at least 17 different ~ tubulins can exist in mammalian
brain MTs (Lee et at., 1986). Tubulin in MTs can also be modifi ed by binding vari ous
li gands. MAPs, enzymati c additi on or removal of amino ac ids (' post- translati onal modi-
fi cati on') and conformati onal state changes. Thus, the number of di fferent possibl e
combinati ons oftubulin states (and infonnat ion capacity) within MTs is extremely large.
In additi on to providing structural support and transport, a role as the ce ll 's nervous
system is suggested by evidence that links the cytoskeleton with cogniti ve functi on. For
exampl e, producti on of tubulin and MT acti viti es correlate with peak learning, memory
and experi ence in baby chi ck brains (Mil eusni e et at., 1980) . When baby rats first open
their eyes, neurons in their visual cortex begin producing vast quantiti es of tubulin
(Cronley-Dill on et at., 1974) . Selecti ve destructi on of animal brain MTs by the drug
colchi cine causes defects in learning and memory whi ch mimi c the sympt oms of
110 S.R. HAMEROFF
Alzheimer 's di sease (i n whi eh neuronal eytoskeleton beeomes tangled and dysfunetional
(Bensimon and Chernat, 199 1) ). 16
The cytoskeleton could serve cogniti ve functions by propagation, storage and process-
ing of informat ion within neurons and other cell s. Tubulin subunits in closely arrayed
MTs have a density of about 10
17
per em', elose to the theoretieal limit for eharge
separation and thus have maximal density for informat ion representat ion by charge. In
general, proteins function by coupling their conformat ional state to a variety of factors;
in essence each protein is a computer element with multiple inputs determining a specific
confonnat ional state as output.
n
Protein Conformational Dynamics
Proteins undergo confonnat ional motions over a wide range of time and energy scales.
However signifi cant conformat ional changes related to protein funct ion generall y occur
in the nanoseeond (10-
9
see) to 10 pieoseeond (10-
11
see) time seale (Karplus and
MeCammon, 1983). Related to eooperat ive movements of protein sub-regions and eharge
redistributions, these changes are linked to protein function (signal transduction, ion
ehannel opening, enzyme aet ion ete.) and may be triggered by faetors ineluding phos-
phorylation, ATP or GTP hydrolysis, ion fluxes, eleetrie fields, li gand binding, and
neighbouring protein confonnat ional changes. In the case of tubulin within MTs, such
programmable and adaptable states can represent and propagate informat ion.
Frohli eh ( 1968, 1970, 1975) proposed that protein eonformat ional ehanges are eoupl ed
to charge redi stributions such as dipole oscill at ions or electron movements within spe-
eifie hydrophobic protein regions (Figure 8). Hydrophobie regions within proteins are
comprised of non-polar side chains of amino acids whi ch exclude water; general anaes-
thetic gas molecul es apparentl y act there to prevent protein confonnat ional responsive-
ness. Frohlich predicted that quantum-level events such as the movement of an electron
within these hydrophobie regions (for example among resonant bond orbitals of amino
acid side chains like the aromat ic rings of tyrosine, tryptophan or phenylalanine) act as a
trigger or switch for the confonnat ional state of the entire protein. Frohli ch further
proposed that a set of proteins connected in a common electromagnet ic field such as
within a polari zed membrane (or polar polymer eleetret like a mierotubule) would be
exeited coherently ifbi oehemieal energy sueh as protein phosphorylation or ATP or GTP
hydrolysis were suppli ed. Coherent exeitat ion times on the order of 10-
9
to 10-
11
see (the
time domain for funct ional protein confonnat ional changes, and in the microwave or
16 Other evidence for cytoskeletal involvement in cogniti on comes from studies of long term
potentiation ('LTP'), a fonn of synaptic plasticity that serves as a model for leaming and memory
in mammalian hippocampal cortex. MAP2 (found only in dendrites) is linked to membrane receptors
and their activiti es by 'second messengers' (G proteins, cycl ic AMP, phosphatidylinositol, calcium
ions, and protein kinases and phosphatases). MAP2 crosslinks MTs, is essential for LTP and
consumes a large proportion of brain biochemi cal energy by phosphorylation/dephosphorylation
(e .g. Theurkauf and Vall ee, 1983; Wang and Resn ick, 199 1; Surridge and Bums, 1992). Excitatory
neurotransmitter activation of LTP post- synaptic receptors induces rapid dephosphorylation of
dendritic MAP2 (Halpain and Greengard, 1990), causes rearrangement of MAP2 connections on
MTs (Bigot and Hunt, 1990) and potentiates excitatory synaptic pathways in rat hippocampus
(Montoro et aI., 1993). In cat visual cortex MAP2 is dephosphorylated when visual stimulation
occurs (Aoki and Siekevitz, 1985) . In an imals whose bra ins are temporari Iy deprived of oxygen, the
degree of cogniti ve damage correlates with reduction in dendritic MAP2 (Kudo et aI., 1990) . Studi es
of protein kinase C suggest learning and memory are structurall y represented in a more complex and
interconnected sub- synaptic cytoskeleton (Friedrich, 1990) .
QUANTUM COHERENCE IN MICROTUBULES III
\
Figure 8.
Two states of protein (e .g.
tubulin) in whi ch quantum
event within hydrophobic re-
g ion couples to confor ma-
tional change with 10-
9
t o
1O-
11
sec transiti ons. Quantum
even t s hown w ithin
hydrophobic region as elec-
tron (e- ) density probability
topography. Genera l anaes-
th eti cs bind within th ese
hydrophobic regions and pre-
vent confor mati ona l trans-
iti ons by van der Waals forces
acting on quantum event.
gigaHz (GHz) speetral region) were dedueed by Frohli eh, who tenned them aeousto-
confonnat ional transiti ons, or coherent ' pumped' phonons. Such a system of coherent
excitat ions in quantum-mechanical terms is call ed a 'Bose-Einstein condensate ' (Wu and
Austin, 1978). Frohli eh also predieted metastable states (longer-lived eonfonnat ional
state pattems stabilized by loeal faetors) and polarization waves (trave lling regions of
dipole-eoupled eonformat ions).
Experimental evidence for Frohlich excitat ions in biological systems includes obser-
vat ion of GHz-range phonons in proteins (Genberg et al. , 199 1), sharp-resonant non-
thennal effeets of mierowave (GHz) irradiat ion on li ving ee ll s (Grundl er and Keilmann,
1983), GHz- induced act ivat ion of microtubule pinocytosis in rat brain (Neubauer et aI. ,
1990) and Raman deteetion of Frohlieh frequeney energy (Genze l et al. , 1983). Vassil ev
et al. ( 1985) demonstrated propagating signal s in mierotubules.
III
Models of Microtubule Information Processing
Considerat ion of MTs as latt iee arrays of eoupl ed (tubulin) dipoles whi eh eooperat ively
interact with their immediate neighbours leads to two types of systems suitable for
infonnat ion processing: spin glasses and cellul ar automata.
In general, three types of arrangements of dipol es in latt iees may oeeur: a) random, b)
ferroe leetrie (parall el-ali gned) and e) spin-glass (regions of loeall y frozen orientat ions).
Tuszynski et al. ( 1994) have examined behaviour of MTs in thi s eontext; depending on
the values of assumed parameters, MT dipole latt ices may exhibit 'frustrat ion', confli ct
in sati sfying all dipole eouplings. Thi s phase strueture is defined as a spi n- glass whi eh
exists close to the edge of chaos and has properties suitable for efficient information
proeessing and eomputat ion. By sli ghtl y alteri ng temperature and extemal field (both
within physiological conditi ons), MT dipol e coupling latt ices may assume a ferroelectric
phase with long-range order and ali gnment with eapabiliti es to propagate kink-like
excitat ions or solitons. The ferroelectric phase appears to be optimal for propagation of
signals and regulation of assembly/di sassembly.
Coupling among tubulin dipoles within MTs in the eontext of eoherent Frohli eh
excitat ions has been proposed as a bas is for informat ion processing, signalling and
cogniti on in a paradigm of cellular automata, a uni versal system of computat ion in a
latt ice structure. Von Neumann described ce llul ar automata (whi ch include computers as
special cases) as systems that consist of a large number of identi cal 'cell s' connected in
112
Glider
, , ,
. :e'
, I .
~ j e j e ,
,! ... ; .. ;
Time 0
.' '.'
, I!.'
: ~ ' ;
Time 1
S.R. HAMEROFF
Time 2 Time 3
Figure 9. Five time steps fr om Conway' s ' Game of Life' cellul ar automaton. Bl ack dot: ' ali ve' ,
white space: ' dead' . Squares ('cell s' ) with two or three li ve neighbours are ' born' in the next
generati on; cell s with zero or one (insuffi cient nouri shment) or four to eight (overcrowding) di e.
Aft er four time steps, the unchanged glider has moved ri ghtward and downward.
a uniform pattern. The essenti al features of cellul ar automata are: I) at a gi ven time, each
ccll is in onc of a numbcr of finit c statcs (usuall y two for simpli city). 2) Thc cell s arc
organi zcd according to a fixed geometry. 3) Each cell communicates only with other cell s
in its neighbourhood; the size and shape of the neighbourhood arc the same for all cell s.
4) There is a uni versal clock. Each cell may change to a new state at each tick of the clock
(or ' generation' ) depending on its present state and those of its neighbours. The neigh-
bour ' transition' rul es for changing states, though simpl e, can lead to compl ex, dynamic
patterns manifesting chaos, fr actal dimensions, parti al differenti al equations and compu-
tation. Patterns which move through the lattice unchanged are call ed ' gliders' ; Von
Neumann proved mathemati cally that with suffici ent time and cellul ar automaton space,
gliders can sol ve virtuall y any probl em. One popul ar exampl e of cellul ar automata is
Conway's the 'Game of Life' in whi ch each 'cell ' in a rectangul ar grid is ali ve (' on ' ) or
dead (' off' ). Each cell 's state at a gi ven time depends on the state of its eight neighbours
in the previous time step. Cell s with 0-1 (too few) or 4-8 (too many) li ve neighbours will
be ' dead'; cell s with 2 or 3 li ve neighbours (, parents') will be 'ali ve '. Despite these
simpl e rul es, compl ex 'life-like' behaviours can ensue (Figure 9).
Cellul ar automata are theoreticall y advant ageous for mol ecul ar computing because the
internal connections are intrinsic to the materi al, external connections need onl y occur in
one limited region, and computation can occur by local interactions with speed dependent
on the clocking fr equency. Quantum automata, in which each state can exi st in 1,0 or
ali ve/dead superposition (like Schrodinger 's cat) have been proposed (e.g. Albert, 1983).
An extreme view has been taken by Fredkin (e.g. Wright, 1985) who contends that the
uni verse is a cellul ar automaton - a latti ce of interacting sub-atomi c logic units, each
one deciding billions of times per second whether it will be ' off' or ' on' at the next
instant. Conrad (1974) used the concept of ' mol ecul ar ' automata within neurons as an
infonnation processing system subserving the brain 's synapti c connectioni sm. In a seri es
of papers (e.g. Hameroff et al., 1982; 1984; 1986; 1987; 1989; 1992; Rasmussen et al.,
1990) cellul ar (' mol ecul ar') automata principl es were appli ed to the dynamic conforma-
tional states oftubulin within cytoskeletal microtubul es (,microtubul e automata' ; MTA).
A rough estimate for the time steps, assuming one coherent phonon or 'sound' wave
across the MT di ameter ::25nm and V sound :: I 0
3
meters/second, yi elds a clocking fre-
quency of approximately 4x I 0" Hz and a time step of2.5x I 0- " second. Thus Frohlich's
coherent excitations can provide a ' clocking fr equency' for MT automata. Transition
rul es were deri ved for each tubulin using electrostatic coupling forces from each of its
six neighbours (Figure 10).
QUANTUM COHERENCE IN MI CROTUBULES 11 3
N
Figure 10 Mi crotubul e automata (MTA) neighbourhood: left, definiti on of neighbourhood dimers;
centre, and monomers within each dimer; ri ght, di stances in nm and ori entation among latti ce
neighbours. Transiti on rul es for MTA are based on net electrostati c coupling forces on each tubulin
fr om its surrounding six tubulin neighbours. Coupling force va lues (described in text) are given in
Hameroff et aI. , 1989 and Rasmussen et aI. , 1990.
The net foree
2 6
e Yi
! ller=-4 L 3
1t ,..
i = I i
where y and r are defined as illustrated in Figure 10, e is the electron charge and E is the
average protein pcnniti vity. yi elds forces of 1.5 picoNcwtons. Energy per tubulin state
change is estimated as 10-
21
Joul es, an order of magnitude greater than random thermal
fluctuations without considering cooperati ve, shi elding effect s from MT-ordering or
surrounding water.
Using these transition rul es, simul ations of MT automata reveal gliders (including
bidirectional gliders), travelling and standing wave patterns, oscill ators, linearl y growing
patterns, and frozen patterns (Figure II). What funetions eould sueh gliders and patterns
have? They eould represent infonnation being transmitted through the eell ; glider
numerical quantiti es and patterns may manifest signals, binding sites for ligands, MAPs
or material to be transported. By determining MAP binding sit es , MT-assembl yl
di sassembly. cytoskeletal network architecture and thus cell structure and function could
be regul ated. Frozen patterns may store infonnation in a memory context, information
may become ' hardened' in MT by post-trans lational modifications, and/or MT automata
patterns could transfer and retri eve stored infonnation to and from neurofil aments via
MAPs.
MT automata gliders trave l one dimer length (8 nm) per time step (10-
9
to 10-
11
see)
for a ve loeity range of 8 to 800 meters per seeond, eonsistent with the speed of trave lling
nerve-membrane potenti als. Thus travelling MT automata patterns or gliders (equi valent
to phonons, Frohlich depolari zation waves, kink-like excitations or solitons) may propa-
gate in the cytoskeleton in concert with membrane depol ari zations and ion fluxes.
Membrane-related voltages, ion fluxes or direct structural links could induce transient
waves of eonfonnational switehing along parall el-arrayed MT. Consequently, external
inputs and acti vity of a particul ar cell could directl y elaborate patterns within that cell ' s
114 S.R. HAMEROFF
1 .. . ..
.. " ..
<x<
2
<:fo; ...................... . .
3
4
:.",",', ',':.',.
.. i)';.(
... .............. y: ..
. .
{n:;
............... ( (.(.( ..... .
.......... ...... . . . -J .. .
..... . . . .. .
.................................. .
H>:< . .>: :
7 ~ " " " .. . .
. . . . . . . . > <X H<' .......... .
: : ~ : : : : : ~ : : : : . '

... . ....
....................... .
.... ( ... ( ... ( ... ( .. ..
.. . . . .. .
.. .. , ..
.............. .
~ ' . ' .. "'" ................. . .... .;.
~ ... " ... ""'''''''''''''''' ........... ~ ..
Figure 11. Eight time steps ofa mi crotubule automata (MTA) computer simul ation. MT is shown
as horizontal open rectangle. Initi al pattern is black 'seeds' (beta state conformations oftubulin) on
wh ite background (alpha state confonn ations) . Seeds 'wiggle' , expand bidirectionally and organize
into ' bus' gliders wh ich move leftward at a veloc ity of8 to 800 meters per second (from Hameroff
et al . 1989) .
MT automata - phenomena important in cogniti ve and behavioural functions ranging
from simpl e organisms to human brains. Long-range cooperat ivity, resonances and phase
transitions emerging from locall y-detennined mi crotubul e automata patterns may occur
among spatially-distributed microtubul es as quantum coherence phenomena and corre-
late with some aspect of consciousness.
Vi ewing the cytoskeleton as the cell 's nervous system, with microtubules serving as
processors or signal carriers and MAPs as mol ecul ar cross-bridge connect ions ('syn-
apses ' ) suggests that the cytoskeleton within each of the brain 's neurons could be viewed
as a 'fractal-like' sub-dimension in a hi erarchy of adapt ive networks. Thi s is consistent
with Globus's (1992) observation that ' nervous ti ssue has fractal geometric, stati stical ,
and dynamic propert ies and that there are fractal correlates of cogniti ve act ivity' . Explor-
ing only a few facets of the possibl e parameter space for cytoskeletal automata, an
enonnous information-processing capacity is impli ed. For example, the 'strong AI'
informat ion-processi ng capacity ofa human brain based on considerat ion of conventional
neural synapt ic transmi ssions as fundamental switches has been estimated as 100 billion
neurons, each with over 1000 synapses 'switching' 100 times per second to yield
approximate ly 10
16
' bits' per second (Moravec, 1987). Consideri ng the simplest case of
microtubul e automata (two confonnat ional states per microtubul e subunit switching at
10-
9
sec. intervals, 10
4
microtubul e subunits per neuron) yields about 10
23
brain ' bits '
per second.
Whether computer-like information processing and molecular automata in the cyto-
skeleton can help explai n consciousness remains questionabl e. Considering only local
interactions, the concept takes the reductioni st argument to a mol ecular level, increasing
the complexity of the brain (and goal for strong AI) about 7 orders of magnitude. but does
not address emergent properti es of consciousness. For thi s, quantum coherence provides
a possibl e resolution.
QUANTUM COHERENCE IN MICROTUBULES 11 5
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