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IRELAND BioSystems 32 (1994) 195-209

Quantum optical coherence in cytoskeletal microtubules:


implications for brain function

Mari Jibu *a'b, Scott Hagan c, Stuart R. Hameroff ~, Karl H. Pribram e, Kunio Yasue ~
"Department of Anesthesiology. Okayama University Medical School, Okayama 700, Japan
hResearch Institute for lnformatics and Science, Notre Dame Seishin University, Okayama 700, Japan
CDepartment of Physics, McGill University, Montrdal, Qudbec, H3A 2T8 Canada
dDepartment of Anesthesiology, College of Medicine, University of Arizona, Tucson, AZ 85724 USA
eCenter for Brain Research and Informational Sciences, Radford University, Radford, VA 24142, USA
f Research Institute for lnformatics and Science, Notre Dame Seishin University, Okayama 700, Japan

(Received 10 August 1993)

Abstract

'Laser-like,' long-range coherent quantum phenomena may occur biologically within cytoskeletal microtubules. This
paper presents a theoretical prediction of the occurrence in biological media of the phenomena which we term
'superradiance' and 'self-induced transparency'. Interactions between the electric dipole field of water molecules con-
fined within the hollow core of microtubules and the quantized electromagnetic radiation field are considered, and
microtubules are theorized to play the roles of non-linear coherent optical devices. Superradiance is a specific quantum
mechanical ordering phenomenon with characteristic times much shorter than those of thermal interaction. Conse-
quently, optical signalling (and computation) in microtubules would be free from both thermal noise and loss. Superra-
diant optical computing in networks of microtubules and other cytoskeletal structures may provide a basis for
biomolecular cognition and a substrate for consciousness.

Key words: Quantum theory; Quantum coherence; Photon coherence; Neural holography; Microtubules; Spontaneous
symmetry breaking; Water molecules; Photon signalling network; Consciousness

1. Introduction presumably patterns composed of many states of


many entities, are distributed throughout the
Several features of brain activity have prompted brain. Despite this extension in space, con-
connections to quantum physics. One feature is sciousness has a 'unity' that is difficult to explain
that the physical correlates of consciousness, by classical means (Marshall, 1989). Non-local
quantum coherence may provide the unity
* Corresponding author, Research Institute for Informatics necessary to break the impasse known as the 'bind-
and Science, Notre Dame Seishin University, Okayama 700, ing problem' in neurophysiological terms (Crick
Japan. and Koch, 1990; Singer, 1993).

0303-2647/94/$07.00 © 1994 Elsevier Science Ireland Ltd. All rights reserved.


SSDI 0303-2647(9!1)01430-2
196 M. Jibu et al. / BioSystems 32 (1994) 195-209

Another feature, as argued by Penrose (1989), is A major interest confronting brain science is,
that many brain functions are non-algorithmic and therefore, to probe the possible mechanisms that
non-computational, and thus inexplicable by serial lead to the processing of quanta of information by
and/or parallel neural processing. Quantum the brain.
coherence (in microtubules specifically, according A specific quantum field theoretical approach
to Penrose (1994)) can account for these functions. was originated by Ricciardi and Umezawa (1967).
Other links between consciousness and quantum Without reference to the structural constituents of
theory include probabilistic synaptic functions brain cells, they formulated a relationship between
(Eccles, 1986; Beck and Eccles, 1992), holography quantum physics and memory. They emphasized
(Pribram, 1966, 1971, 1991; Gabor, 1968), and that each neuron specifies a spatially distributed
general application of the uncertainty principle system with quantum mechanical degrees of
(Bohm, 1951; Bohr, 1958). freedom whose physical properties can be
In the mid 1960s the advent of optical holo- understood by quantum field theory. Quantum
graphy heralded a new departure in understanding field theory is a framework of quantum physics
the relationship between brain, memory and per- capable of describing fundamental processes of
ception. The resistance of aspects of memory and elementary particle physics as well as condensed
of perception to relatively extensive brain damage matter physics (Umezawa, 1993). Ricciardi and
indicated that memory storage and perceptual pro- Umezawa (1967) suggested that memory is a quan-
cessing are distributed procedures. Before the tum vacuum state that violates an original
engineering instantiations of Gabor's holographic dynamical symmetry property of the full comple-
equations were available, it was difficult to ment of the spatially-distributed quantum me-
imagine what such a distributed procedure would chanical degrees of freedom in each brain cell.
be like. The introduction of quantum theory in brain
Over the next decades computer programs science is perhaps analogous to the recent advent
inspired by holography were developed that of the new field, quantum cosmology. The fun-
reflected some of the associative characteristics of damental equations of cosmology, recast as quan-
the parallel distributed processes that constitute tum theoretical operator expressions acting on a
the essence of holographic optical procedures. 'wavefunction of the universe' permit an inter-
These 'connectionist', 'neural network' procedures pretation in which the universe is born in
made it possible to stimulate in vitro many percep- imaginary time. A quantum tunnelling process
tual and memory processes and to explore the would then allow the universe to evolve into real
extent and limitations of the simulations. time, according to a proposal by Hawking.
Meanwhile, it was recognised that an un- Similarly, the model advanced by Ricciardi and
modified holographic metaphor was an inappro- Umezawa (1967) relies on the quantum theoretical
priate model for brain processing because its concept of a vacuum state which violates a
spread function is unlimited (infinite). Data from dynamical symmetry property. While the marriage
neurophysiological experiments showed that the of quantum theory and cosmology is uneasy so
cortical function modelled by holography is better long as quantum theory remains uninterpretable
represented by a sinusoid limited by a Gaussian outside the context of a classical environment, the
envelope - - a mathematical formulation put for- union of brain science with quantum concepts is
ward by Gabor (1946) to measure the maximum consistent with currently accepted interpretations.
efficiency with which a telephone message could be Memory of an external stimulus is imprinted as an
sent across the Atlantic cable. Gabor (1948, 1968) ordering of the vacuum state (i.e. the lowest energy
used the same equations used by Heisenberg to state or the ground state). If this ordered vacuum
describe units in microphysics and, therefore, call- state no longer manifests the original dynamical
ed the unit of communication a 'quantum of infor- symmetry property, the Nambu-Goldstone theo-
mation'. rem suggests creation of long-range correlation
M. Jibu et al./ BioSys~ems 32 (1994) 195-209 197

waves with zero energy requirement which partici- mode considered as a Fr6hlich wave propagating
pate in the system's quantum dynamics (Ricciardi in a one-dimensional medium. Davydov described
and Umezawa, 1967). dipolar solitary waves propagating along alpha-
Such a long-range correlation wave can be seen helices within proteins. This mode of solitary wave
as a Bose quantum ('Goldstone boson' or propagation, known in quantum field theory to
'massless boson'). Ricciardi and Umezawa were carry energy and information without loss due to
perhaps the first to suggest the existence of a thermalization, is called the Davydov soliton or
spatially distributed system with the full range of dipolar soliton (Davydov, 1979).
quantum mechanical degrees of freedom in brain About 10 years after the original idea of 1967,
cells, and also introduced quantum ordering and Umezawa together with two colleagues, Takahashi
spontaneous symmetry breaking as tools to and Stuart, proposed a refined physical model of
investigate the biological system. Slightly later, a spatially-distributed system with fully quantum
Fr6hlich (1968) proposed a similar idea for bio- mechanical degrees of freedom, not only within
logical cells in general but with more explicit each brain cell but also among brain cells (Stuart
materialistic formulation. Focusing on a thin layer et al., 1978, 1979). The system extended among
just beneath the cell membrane, Fr6hlich theorized brain cells is thought to be composed of two kinds
that energy can be stored without thermal loss in of spatially distributed degrees of freedom [ubject
this two-dimensional region in coherent, dipolar to quantum dynamics. As a consequence of spon-
propagating waves. Biological molecules with taneous symmetry breaking, these two modes are
dipolar vibrational activity adjacent to cell mem- distinguished as the 'corticon' field and an
branes may thus manifest a globally coherent exchange boson field which interact as a quantum
mode of dynamics so that the thin layer could be dynamical system - - a theoretical model of non-
seen as a biological superconducting medium or a local memory storage and recall processes. More
biological plasma effectively isolated from thermal recently, Jibu and Yasue presented a physically
environments. Fr6hlich waves are predicted to ap- realistic picture of the system of corticon and
pear in a frequency region between l0 I1 and 1012 exchange boson fields, and proposed to call the
s -l called the Fr6hlich frequency. It can be con- new quantum field theoretical framework 'Quan-
cluded that energy supplied to biological cells with tum Brain Dynamics' ('QBD') (Jibu and Yasue,
magnitude equal to the 'Fr6hlich frequency' times 1992a,b, 1993a,b). They describe corticon and
the Planck constant may not be completely ther- exchange boson fields as dipolar vibrational fields
malized but stored in a highly ordered fashion. distributed along protein filaments of the cyto-
Experimental evidence for Fr6hlich excitations skeleton and extracellular matrices and associated
in biological systems include observation of GHz- water fields. Marshall (1989) proposed that
range phonons in proteins (Genberg et al., 1991), Fr6hlich's 'pumped phonons' result in a Bose-
sharp-resonant non-thermal effects of microwave Einstein condensate in brain which is the substrate
irradiation on living cells (Grundler and for consciousness.
Keilmann, 1983), GHz-induced activation of Inspired by application of quantum theoretical
microtubule pinocytosis in rat brain (Neubauer et methods to brain and other biological cells
al., 1990), and resonance Raman detection of initiated by Ricciardi and Umezawa and by
Fr6hlich frequency energy (Genzel et al., 1976; Fr6hlich in the 1960s, other scientists began to
Webb et al., 1977; Webb and Stoneham, 1977; view brain functioning from the microscopic view
Webb, 1980). of quantum physics during the 1980s. Several
Fr6hlich wave,s need not be restricted to thin groups focused especially on networks of filamen-
layers adjacent to cell membranes. Dipolar oscilla- tous proteins which dynamically organize the in-
tions maintained by hydrogen bonds and non- teriors of living cells: the cytoskeleton (Hameroff
localized electrons trapped in hydrophobic regions and Watt, 1982; Del Giudice et al., 1983). Among
of protein molecules may manifest a collective cytoskeletal filamentous polymers (actin, interme-
198 M. Jibu et al,/ BioSystems 32 (1994) 195-209

diate filaments, etc.), microtubules are the most


central to cellular organization and information
processing. Consisting of skewed hexagonal lat-
tices of subunit protein dimers called tubulin,
microtubules self-assemble into hollow cylinders
25 nm across, adapt, transport and govern cellular !"1111
activities. Their hollow geometry and periodic lat-
tice structure have suggested several quantum field
aspects. Hameroff (1987) applied Fr6hlich's
coherent excitations to tubulin subunits in
microtubules as the basis of computational cellular
automata. Rasmussen et al. (1990) showed that
networks of such microtubule automata are
capable of learning. Considering the layer of
ordered water outside and inside microtubules,
Del Giudice et al. (1986) proposed that the forma-
tion of microtubules' cylindrical structure from
°tubulin' subunits may be understood by the con-
cept of self-focussing of electromagnetic energy by
ordered water. Like the Meissner effect for super-
conducting media, electromagnetic energy would Fig. 2. Microtubule (MT) structure from X-ray crystallography
be confined inside filamentous regions around (Amos and Klug, 1974). Tubulin subunits are 8-nm dimers
comprised of ct and 13monomers.
which the tubulin subunits gather. Del Giudice et
al. (1986) showed that this self-focussing should
result in filamentous beams of radius 15 nm,
precisely the inner diameter of microtubules (see
Figs 1-4).

Fig. 1. Electron micrograph of quick frozen, deep, etched neu- Fig. 3. The tubulin dimer protein has two states in which a
ronal microtubules (MTs) and neurofilaments polymerized quantum event (electron mobility, quasi-particle phonon)
with microtubule associated proteins (MAPs). Arrows point at within a hydrophobic region couplesto conformationalchange
neurofilaments; larger diameter structures are microtubules. with 10-9-10-11-S transitions. Cylinders within protein are a-
Scale bar, 100 nm. (With permission from lqirokawa, 1991). helical regions. (Original drawing by Djuro Koruga).
M. Jibu et al./ BioSystems 32 (1994) 195-209 199

sources of either charge carriers or conformational


waves in microtubules suggest a revival of
holographic brain theory of Pribram and others
(Pribram, 1966, 1971, 1991; Gabor, 1968;
Margenau, 1984; Eccles, 1986; Marshal, 1989).
Our current view is that coherence manifests at the
molecular, quantum level rather than (or in addi-
tion to) coherence at the level of neural firing
(Singer, 1993).
Hameroff suggested in very general terms two
key concepts in understanding cytoskeletal brain
activity from the point of view of quantum
physics: microtubules acting as waveguides for
photons and as holographic information pro-
cessors. Holography is an optical phenomenon re-
lated to interference of coherent electromagnetic
waves or photons. The microtubule may be seen as
a waveguide for photons, and Frrhlich's theory
Fig. 4. Schematic drawing of a synapse showing underlying suggests coherent excitations in microtubules. Fur-
cytoskeletal structure,,;. Right: axon terminal with MT-MAP ther, the periodic lattice structure of microtubules
network connecting w!ith synapsin filaments involved in release
may provide periodically arrayed 'slits' (spaces be-
of neurotransmitter vesicles (circles) into synaptic cleft. Left:
ventrieal dendrite with MT-MAP network, two dendritic spines tween dimers) through which photons may pass.
with cytoskeletal structures connected to receptors in synaptic No proof of coherent photon generation or emis-
membrane. (Adapted from Hirokawa, 1991). sion necessary for optical holography has been
found in microtubules or any microscopic biologi-
cal structure. However, self-focused photons and
Another perspective was proposed by Hameroff self-trapped, non-thermalizing wave entities
(1974) in which microtubules were thought to act should be difficult to detect (see Section 4). In any
like 'dielectric waveguides' for photons, that is, case, no thorough theoretical investigation of
quantum dynamical modes of an electromagnetic these concepts has occurred from the standpoint of
wave. Indeed, living tissue transmits light more physics except the water laser (Del Giudice et al.,
readily than non-living material; in experiments 1988).
with mammalian brains, temporal poles and hip- In the present paper, we propose a new quantum
pocampus manifest the maximum light penetra- theoretical framework suggesting microtubules
tion (Hameroff, 1987). This evidence led Hameroff possess ordered and systematic properties yielding
to consider microtubules as waveguides for cooperative quantum dynamics called superra-
photons. Furthermore, he proposed that cytoplas- diance and self-induced transparency. Specifically,
mic interference of coherent sources from and the quantum dynamical system of water molecules
among multiple microtubules may lead to a and the quantized electromagnetic field confined
holographic information processing mechanisms. inside the hollow microtubule core can manifest a
Holographic patterns stored in cytoplasmic sol-gel specific collective dynamic called superradiance by
states may become 'hardwired' by calcium- which the microtubule can transform any in-
induced assembly of finer components of the coherent, thermal and disordered molecular, elec-
cytoskeleton such as actin, actin-binding proteins tromagnetic or atomic energy into coherent
and the micro-trabecular lattice. Lechleiter et al. photons inside the microtubule. Furthermore, it
(1991) imaged coherent cytoplasmic spiral waves will also be shown that such coherent photons
of calcium induced by acetylcholine binding to created by superradiance penetrate perfectly along
membrane receptors of living cells. Coherent the internal hollow core of the microtubule as if
200 M. Jibu et al./ BioSystems 32 (1994) 195-209

the optical medium inside it were made 'transpar- Their real values would be rMT ~- 12 and IMT -----
ent' by the propagating photons themselves. This 102-103 in nanometers. We denote the spatial
is a quantum theoretical phenomenon called self- region inside the microtubule cylinder by V and re-
induced transparency. strict our discussion to quantum mechanical dyna-
Superradiance and self-induced transparency in mics taking place in this spatial region V.
cytoskeletal microtubules can lead to 'optical' neu- Let us introduce a Cartesian system of coor-
ral holography. Neurons (and other cells) may dinates Oxy~ with the xy-plane attached parallel to
contain microscopic coherent optical supercom- one of the two ends of the microtubule cylinder so
puters with enormous capacity like the fictional that the origin O coincides with the centre of the
but famous HAL 9000 of '2001: A Space Odyssey' end cap. The z-axis lies naturally along the longi-
with holographic memory manipulation (Clarke, tudinal center axis of the microtubule cylinder.
1968). Then, any position in the region V can be labelled
This paper is a result of the long-range coopera- by giving its coordinates r = (x,y,z).
tion among physicists and brain scientists pro- The spatial region V inside the microtubule is
viding several by-products: not empty, but is likely to be filled with water
molecules. Of course, there may be other mole-
1. Superradiance and self-induced transparency cules, although their population is relatively small.
of microtubules may be understood to predict We consider the ideal case in which the existence
theoretically the existence of a peculiar phe- of molecules other than those of water can be
nomenon: weak coherent photon emission neglected. However, below we will consider the
from living matter. more realistic case in which we have impurities
2. Quantum coherence among microtubules among water molecules such as anesthetic
spaced hundreds of micrometers apart (which molecules. It is most likely that the density of
in turn can have quantum coherence with water confined inside the microtubule cylinder re-
other microtubules hundreds of micrometers mains almost constant. Therefore, we may be
away, etc.) can solve the unity and binding allowed to fix the total number of water molecules
problems which plague classical theories of inside-the region V, say N.
consciousness. Let us take a look at a typical water molecule,
3. A source of coherence at the biomolecular say the jth water molecule. Here, j running from 1
level can lead to quantum interference and to N denotes the fictitious number labelling the N
holographic information representation at the water molecules in question. Its position is given
sub-cellular level. by coordinates ~ = (xJ,y, zJ). From a physical
4. General anesthesia may be explained by point of view, a water molecule has a constant
blockade of quantum level events which affect electric dipole, and so it can be seen as a quantum
collective cooperative macro-level events. mechanical spinning top with an electric dipole
moment. The average moment of inertia and elec-
2. Superradiance in microtubules tric dipole moment of a water molecule are
estimated to be I = 2mpd 2 with d ~ 0.82 .A and
Let us look at a typical microtubule in the I.t = 2epP with P ~ 0.2 ,A, respectively. Here, rnp
cytoskeletal structure of brain cells or general bio- denotes the proton mass and ep the proton
logical cells. It is a hollow cylinder about 25 nm in charge.
diameter whose wall is a polymerized array of pro- Due to the electric dipole moment #, the water
tein subunits ('tubulins'). Its length may range molecule interacts strongly with the quantized
from tens of nanometers to micrometers, and electromagnetic field in the spatial region V.
possibly further to meters in nerve axons of large Although the water molecule has many energy
animals. For the purpose of simplifying the physi- eigenstates as a quantum mechanical spinning top
cal formulation, we consider the microtubule as a and so it can exchange energy between the quan-
hollow cylinder with radius rMr and length IMT. tized electromagnetic field in many different
M. Jibu et al./ BioSystems 32 (1994) 195-209 201

values, we restrict our discussion to the most likely us divide the electric field operator into positive
case in which only the two principal energy and negative frequency parts
eigenstates take part in the energy exchange. This
coincides with the conventional two-level approxi- E = E ÷ + E- (3)
mation in describing the energy exchange between
Then, the interaction Hamiltonian of the quantiz-
atoms and the quantized electromagnetic field in
ed electromagnetic field and the totality of water
laser theory.
molecules becomes
Then, one sees immediately that the quantum
dynamics of the jth water molecule can be well N
described by a spin variable s i = 1/2a, where IE-(rJ, t)s j + sJ E+(rJ, t)] (4)
17= (Ox,Oy,•z) and the Ox are Pauli spin matrices j=!
denoting the three components of the angular
momentum for spin 1/2. Let e be the energy differ- where
ence between the two principal energy eigenstates
of the water molecule. Its real value is e = 200
cm -t (Franks, 1972). Then, the Hamiltonian
governing the quantum dynamics of the j t h water The total Hamiltonian governing the quantum
molecule is given by eSJz, and so the total dynamics of the electromagnetic field, the dipolar
Hamiltonian for N water molecules becomes vibrational field of water molecules, and their
interaction is given by
N
HWM = ~ ~ Szi (1)
H = HEM + H W M + HI. (6)
j--i

Two energy eigenvalues of the former Hamilto- Since the spatial region V inside the microtubule
nian are -1/2e and 1/2~ reflecting the fact that only cylinder may be considered as a cavity for the elec-
the two principal energy eigenstates with energy tromagnetic wave, it is convenient to introduce the
difference e are taken into account. normal mode expansion of the electric field
Now, let us consider the quantized electro- operator E = E + + E-, obtaining
magnetic field in the spatial region V. It is conve-
nient to describe the quantized electromagnetic E±(r't) = ~a E~(t)e*i(k .,-w,tl (7)
field in terms of an electric field operator k
E = E(r,t). Let us assume for simplicity that the
electric field is linearly polarized, obtaining Here, ~ok denotes the proper angular frequency of
E = eE, where e is a constant vector of unit length the normal mode with wave vector k. We are main-
pointing in the direction of linear polarization. ly interested in the ordered collective behavior
Then, the quantized electromagnetic field in ques- among the water molecules and the quantized elec-
tion comes to be well described by a scalar electric tromagnetic field in the cavity region V. Let us
field E = E(r,t) governed by the usual Hamiltonian introduce hence collective dynamical variables
S ] ( t ) and S for water molecules by

N
1 [ E2d3 r (2)
s (t) = s (t)e (8)
Heu= j v
j=l

Next, we take the interaction between the quan- and


tized electromagnetic field and the totality of water N
molecules by which they can exchange energy in S= (9)
terms of creation and annihilation of photons. Let j=l
202 M. Jibu et al. / BioSystems 32 (1994) 195-209

Then, the total Hamiltonian (Eq. 6) becomes photons without some pumping mechanism. We
can hardly expect the existence of such a pumping
H = HeM + ES - t~]~..~(E-k S-k + S k4-E k4-) (10) light in the brain. We are, therefore, forced to look
k for another possibility with the total Hamiltonian
(Eq. 10) leading to a different mechanism of
coherent photon emission without pumping light.
It seems worthwhile to note here that this total Fortunately, a physical inspection of the form of
Hamiltonian for the system of N water molecules the total Hamiltonian (Eq. 10) reveals that it
and the quantized electromagnetic field in the manifests a dynamical symmetry property not evi-
region V inside the microtubule cylinder is essen- dent in the ground state and so the resulting quan-
tially of the same form as not only Dicke's tum dynamics is known to involve certain
Hamiltonian for the laser system but also that of long-range order creating phenomena due to spon-
Stuart et al. for Quantum Brain Dynamics (Dicke, taneous symmetry breaking (Stuart et al., 1979).
1954; Stuart et al., 1979). Therefore, it can be ex- The spatial dimension of this long-range order,
pected that each microtubule in the cytoskeletal that is, the coherence length lc can be estimated to
structure of brain cells manifests not only the be inversely proportional to the energy difference
memory printing and recalling mechanism in QBD E, obtaining lc ~- hundreds of micrometers.
but also a laser-like coherent optical activity. The Among those long-range order creating phenome-
former seems not so surprising because Jibu and na we may find a specific one in which the collec-
Yasue (1992a, 1993a,b) developed a physical pic- tive dynamics of the majority of water molecules
ture of QBD in terms of water molecules and pro- inside the microtubule cylinder V can give rise to
tein filaments. Hence, we will not discuss this in cooperative spontaneous emission of photons
the present paper. On the other hand, the latter Without any pumping light. Any incoherent and
seems highly surprising because it may open a disordered energy distribution among the water
completely new picture of the fundamental process molecules due to the macroscopic thermal dyna-
of brain functioning, drastically different from the mics of the polymerized array of protein subunits
conventional one. Namely, in addition to the usual (i.e. tubulins) forming the wall of microtubule cy-
pathway conduction of neural impulses in terms of linder can be gathered collectively into coherent
transmembrane ionic diffusions among nerve cells, and ordered dynamics ready to emit coherent
the brain system may possibly use another but photons cooperatively. This laser-like process of
much more microscopic and elaborated fun- coherent photon emission without pumping light
damental physical process in terms of coherent was first introduced by Dicke (1954) and called
photon emission and transfer in the microtubule superradiance.
just like the optical computer with lasers, optical Let us investigate the superradiance in the
fibers and other optical devices. It is this very sur- microtubule cylinder V starting from the total
prising possibility we are going to discuss Hamiltonian (Eq. 10). We assume for simplicity
throughout the present paper. that only one normal mode with a specific wave
However, it is difficult to look for the possibility vector, say k 0, has a proper angular frequency o~k0
of realizing laser devices in brain cells even though resonating to the energy difference e between the
the microtubule may manifest the same quantum two principal energy eigenstates. Namely we have
dynamical behavior, in terms of water molecules
and the electromagnetic field, as is governed by the = h~k0 (11)
Hamiltonian of a typical laser system. This is
because we need initially to pump up the majority and all the other normal modes are neglected. In
of water molecules to the higher energy eigenstate the conventional laser theory, this is known as a
by certain incoherent but high intensity light. In single mode laser. What we analyse below may
actual artificial laser devices, for example, the ini- hence be considered as a single mode super-
tial pumping is due to xenon flash lamps. In other radiance in the microtubule.
words, the laser system cannot emit coherent Since we have only one normal mode with wave
M. Jibu et al./ BioSystems 32 (1994) 195-209 203

vector k0, we may omit all the wave vector indices This means that a pulse mode of the quantized
of the dynamical variables. Then, the total electromagnetic field in the microtubule cylinder
Hamiltonian (Eq. 10) becomes cavity follows the collective dynamics of water
molecules inside the microtubule. In other words,
H = HEM + e S - # ( E - S - + S+E+). (12) once a collective mode with long-range order is
created in the dynamics of water molecules due to
spontaneous symmetry breaking, coherent emis-
The corresponding Heisenberg equations of
sion of pulse modes of the quantized electro-
motion for the thn~'e collective dynamical vari-
magnetic field - - that is, photons - - follows. This
ables, S and S ±, for water molecules and the two
is the mechanism of superradiance.
variables, E ~, for the quantized electromagnetic
The last question is whether such a collective
field are given by:
mode can be realized in the dynamics of water
molecules starting from incoherent and disordered
dS I I

= -i~-r-(E-S - -S+E +) (13) initial conditions. Notice that such an incoherent


dt h and disordered initial dynamical configuration of
water molecules is due to the interaction l~etween
water molecules and thermally disordered states of
dS ± = 4 - i 2 ~ S E ~ :J- i ~_e_S±
- - (14) the conformational dynamics of tubulins. The
dt h h onset of this collective mode can be seen by
rewriting the three Heisenberg equations for the
and collective variables of water molecules (Eqs. 13
and 14) by substituting Eq. 16. Namely, we have

dE+ = ±i 2re# Sa: (15)


dt hV dS*
= ~ S S "~ 4- i ~ S ± (17)
dt

Because of the short length of the microtubule


c y l i n d e r - / M r ~- 10~"-103 nm - - the pulse mode and
propagating along the microtubule cylinder in the
direction of the z-axis stays in the cavity region V dS
= -f3S+S - (18)
only for a short transit time t M r = IMz/C, where c dt
stands for the speed of light. As this transit time of
the pulse mode is much shorter than the
where
characteristic time of thermal interaction due to
the disordered environment, the system of water
molecules and quantized electromagnetic field in 41re#21Mr
this single mode superradiance is free from ther-
/~ - h2V
mal loss and can be considered as a closed system
well-described by the Heisenberg equations of
motion (Eqs. 13-15). Furthermore, the time These are coupled non-linear differential equa-
derivative of the dynamical variables E + of the tions for non-commuting operators S and S +
quantized electromagnetic field can be approx- subject to certain commutation relations, and it is
imated by E + / I M r in the case of a pulse mode not so easy to find their solutions. However, if we
propagating along the longitudinal axis of the regard these equations as ordinary coupled non-
microtubule. Eq. 15 then yields linear differential equations for classical (i.e. com-
muting) dynamical variables, we can find special
solutions with respect to any incoherent and
E + = 4-i 2 r e # l u r S,~ disordered initial conditions, They are known as
hV (16) semi-classical approximations of the quantum
204 M. Jibu et al. / BioSystems 32 (1994) 195-209

Fig. 5. A schematic representation of the process of superradiance in a microtubule. Each oval without an arrow stands for water
molecule in the lowest rotational energy state. Each oval with an arrow stands for a water molecule in the first excited rotational
energy state. The process is cyclic (a -- b -- c -- d - a - b), and so on. (a) Initial state of the system of water molecules in a
microtubule. Energy gain due to the thermal fluctuation of tubulins increases the number of water molecules in the first excited
rotatinal energy state. (b) A collective mode of the system of water molecules in rotationaUy excited states. A long-range coherence
is achieved inside a microtubule by means of spontaneous symmetry breaking. (c) A collective mode of the system of water molecules
in rotationally excited states loses its energy collectively, and creates coherent photons in the quantized electromagnetic field inside
a microtubule. (d) Water molecules, having lost their first excited rotational energies by superradiance, start again to gain energy
from the thermal fluctuation of tubulins, and the system of water molecules recover the initial state {a).
M. dibu et al. / BioSystems 32 (1994) 195-209 205

dynamical system of superradiance (Agarwal, 3. Self-induced transparency in microtubules


1971). A straightforward calculation yields that
the intensity of coherent photon emission in the We have shown the possibility of a completely
microtubule cylinder due to superradiance can be new mechanism of fundamental brain functioning
given in this approximation by in terms of coherent photon emission by superra-
diance in microtubules. Unlike a laser, superra-
diance is a specific quantum mechanical ordering
hz t - to process with a characteristic time much shorter
I- - - sech 2 (19)
(4tR/~)2 2tR than that of thermal interaction. Therefore,
microtubules may be thought of as ideal optical
encoders providing a physical interface between
where
(1) the conventional macroscopic system of
c h2V
classical, disordered and incoherent neural dyna-
mics in terms of transmembrane ionic diffusions as
tR - 4~rlx2eNiMr
well as thermally perturbed molecular vibrations
and (2) the as yet unknown microscopic optical
and to = tRln2N denotes the life time and delay of computing network system of ordered and
the superradiance, respectively. Notice that the coherent quantum dynamics free from thermal
intensity of superradiance is proportional to N 2 noise and loss. As Feynman (1985) proposed, the
and its delay time is inversely proportional to N. optical computing network is the most realizable
These facts are characteristic to the long-range quantum mechanical computer among many
order creating process involving N water mole- possibilities such as a superconducting computer.
cules (see Fig. 5). However, it is not clear whether the pulse mode
We have found that the quantum collective coherent photons created in the microtubule cy-
dynamics of water molecules and a quantized elec- linder by superradiance can be safely transmitted,
tromagnetic field inside the microtubule cylinder preserving its long-range coherence. It seems most
manifests the long-~rangecooperative phenomenon likely that even coherent photons emitted by
of superradiance in which collective excitation of superradiance will lose immediately their
water molecules can be induced by incoherent and coherence and long-range order due to the noisy
disordered perturbations due to the macroscopic thermal environment. In any case, we absolutely
thermal dynamics of protein molecules forming require some quantum dynamical mechanism to
the wall of the rrdcrotubule cylinder. This fact maintain the coherent transmission of photons in
ensures that each microtubule in the cytoskeletal the as yet unknown microscopic optical computing
structure of brain c,ells, that is, neurons and astro- network in brain cells. In this section, we will show
cytes, may play an important role in the optical that it is again the microtubule in the cytoskeletal
information processing regimen of brain function structure of brain cells which provides us with
as a superradiant device which converts the macro- such a mechanism.
scopic disordered dynamics of water molecules Let us suppose that the pulse mode coherent
and protein molecules into the long-range ordered photons are created in a small segment of the
dynamics of water molecules and a quantized elec- microtubule cylinder by superradiance. Then,
tromagnetic field involving a pulse mode emission those photons propagate along the longitudinal
of coherent pho~tons. In other words, each axis of the microtubule cylinder, that is, the z-axis.
microtubule is a coherent optical encoder in a If the region V inside the microtubule cylinder
dense microscopic optical computing network in were maintained at vacuum, they would transmit
the cytoplasm of each brain cell, if such a network through the region just as they do along a
is realized in actual cytoplasmic structure. This last waveguide. However, the region is filled up with
point is far from evident and deserves to be water molecules, and it is not evident at all
discussed in the following section. whether the pulse mode coherent photons can be
206 M. Jibu et al./ BioSystems 32 (1994) 195-209

safely transmitted through the region without Here,


absorption or loss of coherence.
For the purpose of describing the transmission
of pulse mode coherent photons along the region r=~/2~r~#2N
- ) h 2(t-~
V
inside the microtubule cylinder, it seems conve-
nient to work with a semi-classical approximation
and
in which the long-range ordered dynamics of water
molecules is described by quantum mechanics but
the electromagnetic field is described classically. ~ 27re#2N z
This approximation becomes reliable when the in- ~= h2V c
tensity of the pulse mode coherent photons is
large, and this is indeed the case for superradiance
This is a typical non-linear partial differential
emission (Feynman et al., 1957).
equation called the Sine-Gordon equation, and
Let us consider the Maxwell equation for the
several exact solutions are obtained by means of
scalar electric field E = E(z,t), standing for the
the inverse scattering method of soliton equations
pulse mode coherent photons propagating along
(Ablowitz et al., 1974).
the z-axis coupled to the collective dynamical vari-
The most interesting solution of the Sine-
ables S* of water molecules inside the micro-
Gordon equation (Eq. 23) gives rise to an explicit
tubule cylinder,
form of the time evolution of the scalar electric
field E in the region V inside the microtubule
OE* OE* 2~re# S ~
-- + - :~ i (20) cylinder
Oz #t hV

This equation is valid under the condition that the


_[ 2x~Nvo . [ 2~re#2Nv0 (t z
collective dynamical variables S + are slowly vary- E = "XI V(c - vo)Secn'~ h2V(c - Vo) - ~o) (24)
ing, that is,

OS ± This is nothing but a soliton solution and tells us


- - < < ionS± (21)
Ot that the pulse mode photons propagate along the
dielectric waveguide of the microtubule cylinder
filled up with water molecules with a certain cons-
Then, taking the expectation of those quantum
tant speed v0 less than the speed of light in vac-
mechanical variables in this Maxwell equation and
uum c. It is important to see that the pulse form
the Heisenberg equations of motion (Eqs. 13 and
of the soliton solution is kept unchanged due to
14), eliminating all the expectation values of those
the non-linearity of the Sine-Gordon equation.
variables referring to water molecules, and in-
We have found that microtubules play the role
troducing new variables for the scalar electric field
of dielectric waveguides and that pulse mode
by
coherent photons propagate through them as if
they were perfectly transparent. This phenomenon
2# I}_~.
O±(z,t) = ---~ t E±(z,u)d u (22) is termed self-induced transparency and known to
be a typical non-linear effect in quantum optics
(McCall and Hahn, 1967). The microtubule may
we can obtain dynamical equations for the elec- be an ideal microscopic optical device for use as a
tromagnetic field in the region V inside the perfectly transparent pathway for pulse mode
microtubule cylinder photons, free from thermal noise and loss. Com-
bined with superradiance, this self-induced trans-
020+
- - - sin0* (23) parency of the microtubule allows us to conclude
a~a~ that the brain is essentially a dense assembly of
M. Jibu et al. / BioSystems 32 (1994) 195-209 207

microscopic and e]iaborated optical computing information representation would result from
networks of microtubules in the cytoskeletal struc- quantum optical effects. At the cytoskeletal
ture of brain cells. Coherent photon emission and level, this could occur via calcium induced sol-
transfer in each microtubule are ensured by super- gel variations causing polymerization of actin
radiance and self-induced transparency character- or other cytoskeletal filaments resulting in a
istic to long-range ordering phenomena in holographic representation of short-term
quantum dynamics. memory (Hameroff, 1987). The holographic
Both superradiance and self-induced transpar- paradigm suggests coherence and holographic
ency may be essential to cooperative physical interference across multiple scales (Pribram,
activity of multiple microtubules in cytoskeletal 1991). At a more 'macro' level, coherence of
structure in single cells and among many cells. neuronal firing among widely dispersed neu-
Thus biological systems in general and the brain in rons in the 40-Hz range may be related to
particular may utilise a quantum level holographic quantum coherence of microtubules within
information system (Hameroff, 1987; Pribram, coherently firing neurons (Singer, 1993).
1991). . Sequelae of quantum events in individual
microtubule subunits, acoustoconformational
4. Implications of brain function vibrations could propagate as signals through
cytoskeletal networks and membranes and ex-
We have developed a theoretical framework for tracellular matrices. Such proposed signals (in
the occurrence of quantum coherence of photons the context of the cellular automata theory)
in microtubules and their associated water. Fun- have been shown theoretically capable of
damental involvement of microtubules and the learning, logic and adaptation (Rasmussen et
cytoskeleton in or~zhestration and control of 'real al., 1990; Lahoz-Beltra et al., 1993).
time' cell functions suggests that such quantum
coherence, if it ex!ists, could play a major role in We have described a complex energy/informa-
cellular communication, signalling and informa- tion field originating in cytoskeleton and associ-
tion processing (Conrad, 1988). ated water and cooperatively pervading other
Several aspects of quantum coherence in micro- biostructures. 'Unifying field theories' have a long
tubules may be considered: and unsuccessful history in biology and biophysics
and connote a derisive term: 'vitalism'. We intro-
1. Superradiance and self-induced transparency duce a neo-vitalistic, quantum coherent holo-
occurring in ordered water within the hollow graphic field based on microtubule-derived
core of cylindrical microtubules behaving as Goldstone bosons as a medium of biological infor-
waveguides will result in coherent photons. mation, communication and representation of
This coherence, estimated to be capable of external world. In its largest 'brain-wide' scale,
superposition of states among microtubules this should constitute consciousness.
spatially distributed over hundreds of micro- Fueled by biochemical energy from ATP and
meters, which in turn are in superposition with GTP hydrolysis and protein phosphorylation,
other microtubules hundreds of micrometers microtubule quantum coherence and self-
away in other directions and so on, could focussing should be sensitive to general anesthesia.
account for a coupling of microtubule dyna- Anesthetic gas molecules reversibly inhibit con-
mics over wide areas. This in turn could sciousness by weak, van der Waals binding in
account for a unity of thought and con- hydrophobic regions of proteins (Franks and Lieb,
sciousness. For example, Insinna (1992) has 1982). If quantum optical coherence in micro-
attributed to quantum coherence in micro- tubules is essential for consciousness, anesthetics
tubules the phenomenon of Jungian syn- must somehow inhibit it. One possible way is by
chronicity. membrane inhibition which could disconnect and
2. Coherent interference leading to holographic isolate cytoskeletal dynamics from the external
208 M. Jibu et al./BioSystems 32 (1994) 195-209

milieu. Alternatively, anesthetics may directly alter is partially supported by NSF Grant No.
cytoskeletal quantum coherence. General anesthe- DMS-9114503.
tics do bind to microtubules and at high enough
concentrations cause depolymerization (Allison 7. References
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