BioSystems 109 (2012) 356366

Contents lists available at SciVerse ScienceDirect

BioSystems
j our nal homepage: www. el sevi er . com/ l ocat e/ bi osyst ems
Electrodynamic eigenmodes in cellular morphology
M. Cifra
Institute of Photonics and Electronics, Academy of Sciences of the Czech Republic, Czech Republic
a r t i c l e i n f o
Article history:
Received 29 March 2012
Received in revised form14 June 2012
Accepted 15 June 2012
Keywords:
Cell
Electromagnetic eigenmodes
Cavity resonator
a b s t r a c t
Eigenmodes of the spherical and ellipsoidal dielectric electromagnetic resonator have been analysed. The
sizes and shape of the resonators have been chosen to represent the shape of the interphase and dividing
animal cell. Electromagnetic modes that have shape exactly suitable for positioning of the sufciently
large organelles in cell (centrosome, nucleus) have been identied. We analysed direction and magnitude
of dielectrophoretic force exerted on large organelles by electric eld of the modes. We found that the
TM
1m1
mode in spherical resonator acts by centripetal force which drags the large organelles which have
higher permittivity than the cytosol to the center of the cell. TM-kind of mode in the ellipsoidal resonator
acts by force on large polarizable organelles in a direction that corresponds to the movement of the
centrosomes (also nucleus) observed during the cell division, i.e. to the foci of the ellipsoidal cell.
Minimal required force (10
16
N), gradient of squared electric eld and corresponding energy (10
16
J)
of the mode have been calculated to have biological signicance within the periods on the order of time
required for cell division. Minimal required energy of the mode, in order to have biological signicance,
can be lower in the case of resonance of organelle with the eld of the cellular resonator mode.
In case of sufcient energy in the biologically relevant mode, electromagnetic eld of the mode will
act as a positioning or steering mechanism for centrosome and nucleus in the cell, thus contribute to the
spatial and dynamical self-organization in biological systems.
2012 Elsevier Ireland Ltd. All rights reserved.
1. Introduction
Morphogenesis is long-standing and intriguing problem in
biology. Based on great amount of experience of embryologists,
developmental biologists and cell biologists in microscope obser-
vations, it is apparent that the shape of the organism, be it single cell
or multicellular embryo, is regulated as a whole (Beloussov, 2008).
Such a regulation requires that the behavior of one spatial region
of the organism is intimately connected to the behavior of other
region. Looking for concepts which could represent physical nature
of such holistic regulation, the concept of eld lends itself directly.
Field concept of certain physical quantity can be simply imagined
in a picture in which quantity has certain magnitude (scalar eld)
or certain magnitude and direction (vector eld)
1
in every point of
space and time. Fields are governed, depending on their type, by
fundamental laws which can be described by mathematical equa-
tions. The nature of the eld, materials as well as material interfaces
(described by boundary conditions) present in spatial domain in
which the eld is active restrict the possible eld shapes.
E-mail address: cifra@ufe.cz
1
Example of the scalar eldis temperature eld, example of vector eldis electric
eld.
Inelectrodynamics andalsoinphysics ingeneral, under theterm
mode, we understand a specic shape of the eld having specic
frequency. Eigenmodes
2
of the structure are set of modes which
can be supported by given structure. Concept of eigenmodes is fas-
cinating when one realizes that mere structure gives the form to
the input energy. In other words, when the structure is excited
with (electromagnetic) energy, it will distribute according to the
geometrical and material properties of the structure into the eigen-
modes. In technical physics, we usually use solid state materials to
support modes of interest. Therefore the shape of the structure is
not inuenced by the energy of the eigenmode in such rigid non
developing (non growing) systems. However, if the systemis struc-
turally dynamic (growing or dynamically reorganizing) it may be
able to respond to the eigenmode shape.
Biological systems have been predicted and observed to gen-
erate electromagnetic eld in multiple frequency bands fromvery
lowfrequencyof fewHzuptotheoptical regionas reviewedbyCifra
et al. (2011). Electrodynamic forces generated by cellular skeleton
have been predicted to play role in morphogenesis by Ku cera and
Havelka (2012). Here we are wondering which shapes can electro-
magnetic eld assume on cellular level based on the knowledge of
electromagnetic and geometric properties of biological cells.
2
eigen fromGerman meaning own, self.
0303-2647/$ see front matter 2012 Elsevier Ireland Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.biosystems.2012.06.003
M. Cifra / BioSystems 109 (2012) 356366 357
Biological cell exhibits rich variety of structures. The idea that
cellular cavity structures could function as waveguides and res-
onators of electromagnetic eldis not new. Actually, therearemany
membrane bound organelles (where membrane could impose nec-
essary boundary conditions) which attracted attention of several
researchers.
Popp and Rattemeyer treated nucleus with DNA as a resonat-
ing structure (Rattemeyer, 1978; Popp et al., 1989; Van Wijk and
Shen, 2005; Poppand Beloussov, 2003) for interpretationof photon
emission frombiological systems (biophoton emission).
Cope (1973) treated mitochondria as coaxial resonators for IR
region. The lipid bilayer membrane could serve as a dielectric
between inner and outer conductor (water regions surrounding
membrane are rather lossy in IR region). It was proposed that the
wave guiding property of mitochondrial membrane could serve
for transport of the energy along mitochondrion by means of IR
photons.
Possible resonances of microtubules were treated by Jelnek and
Pokorn y (2001). Cut-off frequency for the rst waveguide modes of
microtubule lies in soft X-ray region. It was speculated that this
kind of modes could interact with biologically signicant atoms of
microtubules and contribute to their dynamic behavior.
Pietak just recently studied geometrical correlation of selected
electromagnetic eigenmode patterns in leaves (Pietak, 2011a,b)
and developing fruits (Pietak, 2012) with the morphology of those
tissues. Frequencies of the eigenmodes were found approximately
intheEHF regionextremelyhighfrequencies (30300GHz). Strik-
ing geometric correlations of eigenmode shapes with the tissue
morphologywereconsideredastructural evidencefor theroleelec-
tromagnetic eigenmodes in plant and fruit tissue morphogenesis.
In our earlier work (Cifra and Lampa, 2008; Cifra, 2010, 2009),
we proposedthat certaincellular electromagnetic eigenmodes may
play role in positioning of large organelles. We analysed the eigen-
modes of cell under approximation of cell as a cavity with perfectly
conducting boundary and lled with homogeneous, lossless and
isotropic dielectric medium. This analysis enabled us to nd the
shapes of electromagnetic cellular eigenmodes. However, approx-
imation with perfectly conducting boundary did not correspond
with the electric properties of the living cell and did not enable us
to determine the quality factor of the modes in a real cell, because
model of resonator with perfectly conducting boundary and loss-
less mediumleads to innite quality factor.
Intherst part of this paper, weextendour earlier workandpro-
vide here analytical treatment of eigenmodes inspherical dielectric
resonator with lossy and isotropic dielectric medium. Further we
employ COMSOL Multiphysics
3
for the numerical calculation of the
eigenmodes and eld distribution in the ellipsoidal resonator. Ana-
lytical methods used for analysis of the eld in spherical resonator
are standard in classical electrodynamics and described in many
textbooks and papers such as (Balanis, 1989; Gastine et al., 1967;
Julien and Guillon, 1986; Yadav and Singh, 2004). In the second
part of this paper, we analyze the results in terms of contribution
to organization in living cells.
2. Positioning of Centrosome and Nucleus
Centrosome is organelle present in many types of cells and
is responsible for nucleation of microtubule growth. Therefore
centrosome behaves as microtubule organizing center. Since
microtubule cytoskeleton is important for organization of struc-
tures and processes within the cell, the position of centrosome as
microtubule nucleator is crucial for the cell function. There is well
3
www.comsol.com.
known observation that the centrosome always tends to be posi-
tionednear toor inthe center of the nondividing cell andassociated
near the nucleus (Alberts et al., 2008; Euteneuer andSchliwa, 1992)
as depicted in Fig. 1.
A model for the centrosome and nucleus positioning behavior
based on force exerted by microtubule polymerization in conned
space has been proposed to explain the centrosome and nucleus
centering by Reinsch and Gonczy (1998), Holy et al. (1997) and
Daga et al. (2006), yet the exact mechanisms enabling this in vivo
are not fully elucidated.
During the cell division the mitotic spindle, crucial structure for
the correct distribution of the identical copies of chromosomes to
newdaughter cells (Alberts et al., 2008), is formed. In animal cells,
mitotic spindle (assembled frommicrotubules) nucleates fromthe
two divided centrosomes. During the creation of mitotic spindle,
both centrosomes move to foci of the dividing cell. Dividing animal
cell can be roughly approximated by ellipsoid, see Fig. 2.
Here we propose that certain eigenmodes of the cell as cav-
ity electromagnetic resonator play role in positioning of organelles
such as nucleus and centrosomes.
3. Fundamentals of Cavity Resonator Electromagnetics
Physically, followingthree characteristics are crucial inthe anal-
ysis of electromagnetic cavity resonator:

Inner bulk material in the volume of the resonator. This is usually

describedinoptics byindexof refractionp =

r
.
r
is relative
4
permittivity (also termed as dielectric constant) and
r
is rela-
tive permeability, this pair of quantities is preferentially used in
radiofrequency physics. These quantities are frequency depen-
dent ingeneral. c
r
describes interactionof the electric component
of the electromagnetic eld with matter and
r
the interaction of
the magnetic component of the electromagnetic eld with mat-
ter. Both
r
and
r
of the material can be complex numbers. Their
imaginary parts indicate that electromagnetic eld is absorbed
by the medium and transformed to another form of energy (e.g.
motion of electrically charged particles, which, if it is disordered,
is macroscopically observed as heat). The ratio of imaginary and
real part of permittivity is oftenexpressedas tan =

being
imaginary part and

being real part of the permittivity. Higher

tan means higher losses of the material

Boundary of the resonator. Electromagnetic eld obeys bound-

ary conditions at the interface of the two materials (e.g.
metaldielectric, dielectricdielectric). Boundary conditions
stemfromthe fact that certain eld components must to be con-
tinuous (without the jump in the value) across the interface
in order for the eld to be physically real. Boundary conditions
used in calculation signicantly inuence the eld shape in the
resonator and need to be carefully treated in order to describe
physical reality.

Size and shape of the resonator. They both inuence the frequen-
cies and shape of the eld in the resonator.
Above mentioned characteristics, when specied, uniquely
determine set of eigenmodes of the resonator. Eigenmodes of the
spherical and spheroidal resonators we treat in this paper can be
categorized into TE
nml
and TM
nml
modes.
5
4
Relative to the permittivity of the vacuum.
5
TE stands for Transverse Electric (EM eld has electric component transversal
to the radius vector, i.e. Er =0), TM stands for Transverse Magnetic (EM eld has
magnetic component transversal to the radius vector, i.e. Hr =0), subscripts n, m,
and l are found in the eld equations (e.g. Gastine et al., 1967). Basically, the higher
the mode numbers nml, the higher is number of eld maxima and minima along
358 M. Cifra / BioSystems 109 (2012) 356366
Fig. 1. (a) Human osteosarcoma cell xed with methanol and stained with polyclonal antibody to alpha-tubulin (green) and monoclonal antibody TU-30 to gamma-tubulin
(red). DNA is stained by DAPI. (Photography E. Draberova, IMG AS CR, Prague) (b) Schematically depicted location of the centrosome in the interphase of the model spherical
cell. Two perpendicular rectangles are centrioles, which nucleate growth of microtubules in animal cells. Lines represent microtubules. Nucleus is not depicted here.
Fig. 2. (a) Human osteosarcoma cell xed with methanol and stained with polyclonal antibody to alpha-tubulin (green) and monoclonal antibody TU-30 to gamma-tubulin
(red). DNA is stained by DAPI. (Photography E. Draberova, IMG AS CR, Prague) (b) Schematically depicted location of the centrosomes and shape of mitotic spindle during the
division of the cell. Two perpendicular rectangles depict centrioles, which nucleate growth of microtubules in animal cells. Lines represent microtubules.
Eigenmodes inthe resonators have three important parameters:

shape of the eld (given by eld equations)

frequency at which they can be excited

quality factor Q - it is ratio of how much energy can be stored

in the resonator compared to input energy rate, in other words,
ratio of energy stored in the resonator and energy lost per cycle
times the frequency, see Eq. (1).
Quality factor Q for dielectric resonator is dened as
Q =
energy stored
energy lost per cycle
=
W
P
d
+P
r
(1)
where =2f, f being frequency, Wis energy stored in resonator,
P
d
denotes dielectric losses and P
r
denotes losses due to radiation
of energy fromthe resonator.
radial andangular directions. Note that the conventioninliterature is to denote both
principal mode number and index of refraction is denoted as n and both imaginary
part of index of refraction and Boltzmanns constant as k. To avoid the confusion we
will use p to denote index of refraction and q to denote the imaginary part of index
of refraction.
To combine the contribution of losses using corresponding
Q factors, for the dielectric resonator (resonator with dielectric
boundaries) we can also write
1
Q
=
1
Q
d
+
1
Q
r
(2)
Q with subscripts d, r are related to power losses due to the dielec-
tric loss inside the resonator and radiative loss, respectively. For
any resonator and method used, once the frequency f =f

+if

of the
mode is found, it enables one to calculate the quality factor directly
as
Q =
j

2j

(3)
Resonator modes (or any oscillatory process in general) can be
categorized into three regimes based on its Q factor:

overdamped, Q<1/2

critically damped, Q=1/2, energy stored inthe mode is dissipated

into other formapprox. within one period of oscillation

underdamped, Q>1/2.
In the current work, underdamped modes are of our interest,
because they can store energy and thus enable formation of their
eld shape.
M. Cifra / BioSystems 109 (2012) 356366 359
Table 1
Permittivity of the dry biomaterial, water and biomaterial for three frequencies. * - Imaginary part of refractive index q was calculated from absorption coefcient a as
q =a(z/4), where z is free space wavelength, calculated using Eq. (B.1), 75% of material volume being water and 25% dry biomaterial.
Frequency Permittivity
Dry biomaterial Water Biomaterial
1.6THz p=2+i0.2* p=2.15+i0.09*

=3.94+i0.96 Wilmink et al. (2011), Fig. 4 =4.61+i0.41 (muscle tissue,
Wilmink et al. (2011), Fig. 6)
24 THz p=1.52+i0.1 =1.2+i0.55 Ellison
(2007), Fig. 25
=1.44+i0.51

=2.3+i0.23
lyophilized bacterial
cells, Cooper and
Powell (1986), 11m
wavelength (27.3THz),
Fig. 2
50THz p=1.5+i0.1 p=1.25+i0.11 =1.7+i0.29

=2.24+i0.3 see text =1.54+i0.28
(Wieliczka et al., 1989),
1667cm
1
4. Studied Models
We created two model resonator geometries to approximate
interphase nondividing cell (spherical model) and dividing cell
(prolate ellipsoid model). The diameter of size of the model
interphase cell is a =7m and semiaxes of prolate ellipsoid are
a =9.1726m, b=c =6.1151m. Ellipsoidal dividing cell has the
same volume as the spherical interphase cell. Capability of spheri-
cal cells of small (a =3.3m) and large radius (a =65m) to behave
as dielectric resonator is also analysed in order to assess generality
of the model.
4.1. Dielectric Properties of Resonator
Permittivity of any material is generally dependent on fre-
quency. Dielectric properties of the material inside the resonator
(which represents inner volume of the cell) used in our calcula-
tions are based on experimental ndings from measurements of
bulk biological material. The is no single publication which covers
the permittivity data of biomaterial in the whole range of frequen-
cies (wavelengths) of interest (150THz). Furthermore, data from
dry biomaterials only are often presented.
Generally, water is the most abundant component of the bio-
logical cell, therefore it plays the crucial role in the effective
permittivity of the cell. Water is strong absorber of THz radiation
with two major peaks at 6 and 19.5THz (Fig. 5 in Arnone et al.,
2000). Frequency dependence of the permittivity of the water can
be modelled by few relaxation and resonance processes (Ellison,
2007). For more data on water complex permittivity in THz and IR
region, (see Downing and Williams, 1975; Ellison, 2007; Segelstein,
1981; Wieliczka et al., 1989).
In the frequency range where no direct data on naturally wet
biological material are present, theory of mixture of dielectric
materials is used, see Appendix B, assuming that water volume
fraction in cell is 75%. We gathered data on dielectric properties
of water, biomaterial or dry biomaterial and water for the three
frequency ranges: low, middle, high, see Table 1. It was not pos-
sible to obtain the permittivity data for biomaterial in the high
frequency range of interest (50THz, mid IR range) fromthe litera-
ture. Therefore we try to estimate the permittivity values based on
the properties in optical region. It is well known that biomolecules
have higher refractive index than water in optical region. Pro-
tein refractive index is usually in the range of p=1.51.6 (Hand,
1935; McMeekin et al., 1962) while extreme case such as tubulin
has p=2.9 (2002-Mershin-Biosytems). Lipid bilayers have p=1.52
(Ohki, 1968). Carbohydrates have p=1.41.5 (Aas, 1996 Fig. 3).
Based on the dielectric relaxation theory, permittivity at lower
frequencies can assume only higher values, although resonance
behavior may locally perturb this trend. It needs to be noted that
the permittivity in the mid IR region has rich behavior because it
involves frequencies of many molecular vibration modes. Based on
the refractive index and permittivity data in the optical region, we
made an estimation of dielectric properties of biological material
in the mid IR (50THz): p=1.5+0.1i, =2.24+i0.3.
It needs to be noted that the data on permittivity values come
frommacroscopic measurement of biological material. It may rep-
resent some averaged level of losses because eld conguration
used in technical macroscopic measurement is rather uniformand
therefore substantially different from eld conguration in cav-
ity resonator. In such macroscopic measurement, eld sees the
biomaterial oriented rather randomly and that makes it likely to
average out the losses. In the case non-random orientation of the
molecules with respect to the eld on the sub-cellular level losses
could be substantially higher or lower.
4.2. Spherical Dielectric Resonator
For the resonator with dielectric boundary, dielectric properties
of surrounding material are crucially important for the resonator
behavior. We assume that a watery mediumis surrounding the cell
resonator. Water refractive indexandpermittivityhas beenalready
discussed in the previous section.
Using eld equations inside and outside the spherical dielec-
tric resonator (Gastine et al., 1967) and general electromagnetic
boundary conditions (see. e.g. Balanis, 1989) lead to characteristic
equation for TM and TE modes which is both complex and tran-
scendental.
Following characteristic equation is valid for TMmodes
j
n1]2
(k
1
u)
j
n+1]2
(k
1
u)

n
k
1
u
=

H
(2)
n1]2
(k
2
u)
H
(2)
n+1]2
(k
2
u)

1
n

2
k
1
u
(4)
360 M. Cifra / BioSystems 109 (2012) 356366
and for TE modes
j
n1]2
(k
1
u)
j
n+1]2
(k
1
u)
=

H
(2)
n1]2
(k
2
u)
H
(2)
n+1]2
(k
2
u)
(5)
J is Bessel function and H
(2)
is Hankel function of the second kind,
their subscript indices denote the order of the functions. Relative
permittivity of resonator is denoted by
1
, relative permittivity of
the surrounding mediumis denoted as
2
. k
1
=

1
k
0
, k
2
=

2
k
0
,
wave constant k
0
=2f/c
0
, f is frequency, c
0
velocity of light in vac-
uum, a radius of the spherical resonator. Roots of the equations are
complex frequencies which correspond to the frequencies of the
resonator modes. This enables us to calculate the frequency of the
modes and further using Eq. (3) also the quality factor of the modes.
Additionally to analytical calculations, COMSOL Multiphysics
software has been used to calculate the eld and also as a tool for
eld visualization. COMSOL uses Finite Element Method, a numer-
ical method which can be used also for the calculation of the
electromagnetic eld. Eigenmode solver of COMSOL enables cal-
culation of modes of any structure. Since the method is numerical
and the computer memory is limited, there is a limit for number
of points in the space where the eld can be calculated. Therefore,
the domain where the calculation is to be performed needs to be
spatially limited using suitable boundary conditions. We can use
impedance boundary condition if we can nd the correct value of
the impedance of the eld at the boundary. Wave impedance in
spherical coordinates is dened as ratio of perpendicular (to radius
vector) components of electric and magnetic elds. Since elds of
the dielectric resonator are determined analytically by equations
of electric and magnetic eld (found e.g. in Gastine et al., 1967) we
can use following general equation to calculate impedance Z at the
boundary r =a, a being radius of the spherical resonator.
Z =

(6)
where E

is component of the electric eld perpendicular to radius

vector (i.e. 0 or in spherical coordinates) and H

is perpendicular
component of magnetic eld.
However, ingeneral, boundaryimpedance of modes indielectric
resonator differs frommode to mode based ontheir type (TMor TE)
andmode number n. Therefore, whenwe set specic numeric value
of boundary impedance in COMSOL or other numerical software, it
will be valid only for the mode for which it has been calculated.
If other modes are found by the numerical eigenmode solver, they
may not correspondto any other real mode, i.e. they may not repre-
sent any mode which would be present in physically real dielectric
resonator.
4.3. Ellipsoid Resonator
Analytical ellipsoid resonator treatment involves spheroidal
functions (Mathieu functions) and is studied in substantially
smaller amount in the literature (for example by Li et al.,
2002, 2003; Kokkorakis and Roumeliotis, 1997; Mehl, 2009) than
the spherical case. Denition and implementation of general
spheroidal functions is therefore not standard and requires signi-
cant amount of programmingcomputational efforts suchas byVega
(2003) and is beyond the scope of this paper as it is partially active
research topic on its own. However, numerical methods such as
Finite Element Method enable analysis of the eld in any resonator
shapeinprinciple. ThereforeweapplyCOMSOLMultiphysics eigen-
mode solver to calculate the eld in ellipsoid resonator.
Boundary impedance for the mode of dielectric ellipsoid res-
onator was estimated based on the boundary impedance for the
corresponding mode of spherical dielectric resonator radius of
a =8m. This radius was chosen to be between the radius of
Fig. 3. Dependence of quality factor due to radiative losses, Qr, of TM
1m1
mode on
the permittivity contrast INT/EXT.
sphere with volume equivalent to the ellipsoid (a =7m) and the
9.1726m length of the major axis and 6.1151m length of two
minor axes of ellipsoid resonator. Estimated boundary impedance
for the mode of interest was Z =299.96+i44.913!. Whenusing this
method, it needs to be noted that the eld shape and the frequency
of the modes in ellipsoidal resonator with dielectric boundary is
not exact but it describes major geometric features.
5. Cellular Electromagnetic Modes of Biological Interest
Regarding possible biological function, we found the resonator
modes which have shape spatially correlated with cell structures
commonlyobservedunder microscope. We assumethat thepattern
of the eld in the cell determines the occurring structure namely
by attraction of large cellular structures to the region of maxima
of electric eld intensity as will be explained later. For now one
has to keep in mind that we search for modes having maxima of
electric eldintensityinthelocationof centrosomes and/or nucleus
as depicted in Figs. 1 and 2.
5.1. Spherical Resonator Nondividing Cell
Spherical resonator represents a nondividing (interphase) cells.
In the case of dielectric boundary resonator a new classication
of modes additional to TE and TM modes arises compared to res-
onator with perfectly conducting boundary which traps the eld
completely inside the resonator. Modes of the dielectric resonator
can be classied to external and internal modes, as done by Gastine
et al. (1967), due tothe nature of dielectric resonator. It is typical for
external modes to have signicant portionof energy ineldoutside
the dielectric resonator. This is reected in lowquality factor of the
external modes. It may be educative to nd the frequencies and
quality factors of external modes, but the external modes cannot
store much energy (their Q factor is the order of 0.1) due to their
physical nature and therefore are not very interesting regarding
possible biological role.
In the dielectric spherical resonator, the discovered mode of
interest is TM
1m1
. Quality factor due to radiative losses, Q
r
, is purely
dependent onthepermittivitycontrast
INT
/
EXT
, where
INT
is abso-
lute value of permittivity of the resonator dielectric and
EXT
is
absolute value of permittivity outside the resonator. See Fig. 3 for
Q
r
for TM
1m1
for various values of permittivity contrasts. This gure
was obtainedbycalculatingtheQfactor of TM
1m1
fromcomplexfre-
quency obtained from Eq. (4) for various values of permittivities.
The Q
r
values do not depend directly on the resonator diameter.
M. Cifra / BioSystems 109 (2012) 356366 361
For getting the total Q factor of the mode of dielectric resonator,
we can use Eq. (2), where we need to know Q
r
and Q
d
. Once we
know the value of permittivity of the material inside and outside
the resonator, we can nd the Q
r
fromFig. 3 and Q
d
fromEq. (A.2).
This estimation does not take into account the fact that the dielec-
tric losses (tan) in the material inside and outside the resonator
can be different. However, dielectric losses inside and outside the
resonator are of the same order of magnitude in our case and most
of the energy of the eld is stored inside the resonator so no great
deviation from Q
d
estimation based on the losses in the resonator
is expected.
For thespherical cell radius a =7mandpermittivityinthemid-
dle frequency range (24THz), TM
1m1
has frequency of f =24.67THz,
quality factor Q=0.903 (from Eq. (4) and Eq. (3)) and bound-
ary impedance Z =305.04+i72.718! (from Eq. (6)). Calculation
with COMSOL yielded very similar frequency f =24.81THz and
Q=0.9104. The small deviation from analytical value from Eq. (4)
may be caused by the error of numerical method used (FEM nite
element method) which may stem from the nite density of dis-
crete tetrahedral mesh. For the depiction of electric eld of TM
1m1
mode see Fig. 4.
For the small cell with radius a =3.3m), TM
1m1
mode fre-
quency is f =1.67THz and Q=1.3362 and for large radius cell
(a =65m) the TM
1m1
has frequency 44.89THz and Q=0.8737. It
seems that the TM
1m1
modes can be supported within the whole
range of cell sizes. Actually, it is interesting to observe that the res-
onances appear to be feasible starting fromcca. fewTHz, i.e. where
the permittivity ratio
CELL
/
WATER
crosses the unity and rises above
it.
The mode TM
1m1
electric eld distribution in dielectric res-
onator is shown in Fig. 4. Highest intensity of the electric eld is
in the center of the sphere. We are not interested now in absolute
values of the eld intensity since it depends on the excitation. Exci-
tation is normalized for all modes displayed. Therefore we have not
included color bar showing the linear intensity scale. Mode TM
1m1
is triply degenerated.
6
Other two modes with same n and l, but
with different m(not depicted here) are orthogonal to the depicted
mode but with the same resonant frequency. These two degener-
ated modes have maxima of electric eld intensity in the center of
the resonator as well. Actually, external mode TM

1m1
of dielectric
resonator perfectly resembles the shape of TM
1m1
mode of con-
ductive boundary resonator (see e.g. Cifra, 2010, 2009; Cifra and
Lampa, 2008 for study of eld distribution of spherical resonator
with conductive boundary). If we did not accept categorization of
the dielectric resonator modes into internal and external, external
mode TM

1m1
would be denoted as TM
1m1
and the internal mode
we call now TM
1m1
would be denoted as TM
1m2
. The l =2 would
perfectly explain why we observe rise in intensity along the radius
vector after some distance fromcenter because l =2 means we use
second root of the Bessel function, i.e. second local maximumalong
radial directionhas tooccur withinthe resonator volume. However,
this approach to mode renumbering, although useful for under-
standing in this case, could not be used fully generally because
external modes are of different nature thaninternal modes. Namely
frequency and quality factor of external modes, in contrast to inter-
nal modes, are rather weakly dependent on dielectric properties
of inner resonator medium, as explained and comprehensively
described by Gastine et al. (1967).
6
The termdegeneracy means that the modes have same resonant frequency but
different mode shape. This stems fromthe fact that the resonant frequency does not
depend on modal number m, but mdoes inuence shape number and can assume a
certain range of integer values limited by n.
5.2. Ellipsoidal Resonator Model of Dividing Cell
Prolate ellipsoidal resonator represents dividing cell. The mode
of interest is that with highest intensity in the both focal points
of ellipsoid. They have double degeneracy. Distribution of elec-
tric eld intensity in the dielectric boundary resonator is depicted
in Fig. 5. For the ellipsoid of 9.1726m length of the major axis
and 6.1151m length of the minor axes, having permittivity cor-
responding to the middle frequency range (Table 1), the discovered
mode of interest has frequency f =30.108THz and Q=1.5589.
6. Effect of the Electric Field on Organelle Positioning
We theoretically quantify the effect of electric eldof the modes
of interest on cellular morphology in this section. Effect of the eld
onthe matter canbe mediatedby transfer of energy or information.
Here needs to be noted that also information needs to be carried
by some minimumamount of energy. Thus, both transfer of energy
and information involve force of the eld acting on the matter.
6.1. Mechanisms
6.1.1. Electric Field Acting on Charge
Electric eld E can act by force F on charge Q as
F = QE (7)
At high frequencies as those involved in discovered modes of
interest, there will be no net translational force of electric eld
acting on a charge, since the eld is oscillatory.
6.1.2. Electric Field Acting on Dipole
Nonuniformelectric eld can act by translational force on elec-
tric dipolepwhichis commonelectric approximationof manypolar
molecules or macromolecular structures and organelles:
F = (p E) (8)
6.1.3. Electric Field Acting in Induced Dipole Dielectrophoresis
Even if the cellular organelle does not have signicant electric
dipole moment itself, nonuniform electric eld can polarize the
organelle if the organelle permittivity is different from that of the
surroundingmedium, thus creatinganinduceddipolemoment. The
phenomenon of force of nonuniformelectric eld acting on neutral
dielectric particle is called dielectrophoresis and is described thor-
oughly by Pohl (1978). We consider cellular organelles a dielectric
neutral particles whichhavehigher permittivitythanthesurround-
ing cytosol. This can be expected for the high frequencies in the
region of tens of THz calculated for the cellular eigenmodes, since
organelles contain proteins and lipids which have higher optical
permittivity than water.
The dielectrophoretic force is approximately given by
F v(
p

s
)
2
(9)
Fis forceactingonthedielectric particle; Vis volumeof theparticle;

p
is the permittivity of the particle;
s
is the permittivity of the
surrounding media; denotes the gradient; E is electric eld.
The dielectric particle with higher permittivity than its sur-
roundings is forced to move in the direction of gradient of the
squared intensity of electric eld. It is due to the fact that the par-
ticle will have lowest potential energy in the location of highest
intensity of electric eld. Inthe following, we will analyse the effect
of dielectrophoretic force on the organelle positioning.
362 M. Cifra / BioSystems 109 (2012) 356366
Fig. 4. TM
1m1
mode electric eld distribution in dielectric spherical cavity resonator, radius a =7m, f =24.67THz (Q=0.903) (a) plane x =0. White arrows depict electric
eld direction. (b) Quasi 3D viewin three orthogonal cutting planes.
Fig. 5. Electric eld distribution of the TMkind of mode in prolate ellipsoidal dielectric cavity resonator, major axis a =9.1726m, minor axes b=c =6.1151m. f =30.108THz
(Q =1.5589) (a) Plane z =0. White arrows depict electric eld direction (b) Quasi 3D viewin three orthogonal cutting planes.
6.2. Minimal Required Force
In our current study, in the case when the organelle or particle
does not expend its own energy sources for translational move-
ment, the force exerted by electric eld needs to reach certain
threshold above which it will overcome effective force of the
Brownian motion.
7
The threshold force is given by Hlzel et al.
(2005) as
F
th
=

2
D^t
k1 (10)
D is the diffusion constant of the organelle (particle); ^t is time
period of the acting force; k and T are StefanBoltzmann constant
and temperature.
Diffusion constant for rigid sphere (approximation of the
organelle) is given by
D =
k1
viscous drag
=
k1
6qr
(11)
7
Stronger the deterministic force, smaller the deviation of probability distribu-
tion of particle localization fromthe mean position.
q is dynamic viscosity (we take that of water (1mPa s) as approx-
imation) r is radius of the particle (we take 150nm as reasonable
approximationof centrosome radius (Ueda et al., 1999), for nucleus
it would be about 5003000nm)
For the temperature of 293K the threshold force to overcome
Brownian motion is
F
th
= 1.513 10
15
N for^t = 10s (12)
F
th
= 4.783 10
16
N for^t = 100s (13)
F
th
= 1.513 10
16
N for^t = 1000s (14)
The dielectrophoretic force on the spherical particle in inhomo-
geneous electric eld is given exactly by Pohl (1978) and Hlzel
et al. (2005)
F
DP
= 2r
3
Re{
m
CMfactor}
2
(15)
where CMfactor is ClausiusMossotti factor
CMfactor =

p

m

p
+2
m
(16)

p
and
m
being complex permittivity of particle and medium,
respectively.
For example, at 24THz taking
p
=2.3+i0.23 (permittivity
of protein rich particle) and
m
=1.44+i0.51 (permittivity of
M. Cifra / BioSystems 109 (2012) 356366 363
material inside the resonator), we get 2r
3
Re{
m
CM fac-
tor} =4.785610
32
. To reach and overcome the threshold force
for ^t (duration of dielectrophoretic force) of 100s
8
gradient of
squared intensity of electric eld needs to assume the value of
10
16
V
2
m
3
.
6.3. Energy in the Cell
In order to know possible intensity of electric eld and electric
eld gradient of the modes, we rst need to assess the energy avail-
able in the cell. The total thermal power produced by the cell (of
yeast cell type which has typical radius a =4m) was determined
experimentally using calorimetry by Lamprecht (1980) to be about
10
13
W. That means that the energy of about 10
13
J is produced
by cellular chemistry per second. This can be considered an upper
boundary of possible energy production rate in the cell.
Thermal electromagnetic energy based on Plancks law can be
calculated using equation
W
Mthcrm
= b1
4
v (17)
where V is volume and T is temperature in Kelvin.
u = b1
4
=
8
5
k
4
15(hc)
3
1
4
= 7.5502 10
16
1
4
(18)
where u is thermal radiation energy density [J/m
3
], k is
StefanBoltzmanns constant, h is Plancks constant and c is veloc-
ity of light in vacuum. For the cell with radius a =7m, i.e.
volume v =
4
3
u
3
m
3
in room temperature (T =296.15K=23

C)
W
EMtherm
=8.3410
21
J.
Thermal electromagnetic energy W
EMtherm
10
20
J forms a
lower boundary of the range of possible energy available in the
resonator.
6.3.1. Spectral Properties of the Excitation Energy
The most trivial possible case of energy distribution is ther-
mal Planck spectral energy distribution depicted in Fig. 6 based on
equation
u(j )dj = dj
8hj
3
c
3
1
c
hj ]k1
1
(19)
It is interesting to observe that spectral distribution of ther-
mal electromagnetic energy which is omnipresent under common
Earth temperature conditions (037

C) has peak frequency exactly

inthe range of rst electromagnetic resonant modes of cells of com-
mon sizes. In other words, the wavelength of omnipresent thermal
electromagnetic radiation on Earth is just the same as the diame-
ter of most cells (Fig. 6). What could be other possible sources of
electromagnetic eld in the cells for the modes of interest which
lie in frequency region of 150THz (i.e. at the border of middle and
far infrared region)? From molecular point of view, stretch, bend-
ing, rotation of molecular groups lie in this region, as is also the
case in proteins as reviewed by Barth (2007). Compared to other
molecules, proteins are special in that they can convert chemically
bound energy fromphosphate bonds of nucleotides as ATP and GTP
into the other types of energy. IR generation and radiation from
biological systems of above thermal level is poorly explored, partly
due to the experimental difculties, because it can be hard to dis-
tinguish the IR radiation just due to the macroscopic temperature
of the object and due to processes which generate IR nonthermally.
As only work known to the author, Fraser and Frey (1968) detected
8
100s is reasonably short time compared to duration of the cell division which
may take fromtens of minutes to hours
Fig. 6. Spectral distribution of thermal energy, frequency dependence. Frequency
axis is in logarithmic scale with base 10, i.e. 13 correspond to 10
13
Hz = 10THz (free
space wavelength 30m) and 13.5 corresponds to 10
13.5
Hz = 31.6THz (free space
wavelength 10.53m).
IR radiation fromactive crab nerve in the region of 110m. Emis-
sion intensity was 2 orders of magnitude higher than that of the
thermal radiation. This detected intensity of IR radiation is proba-
bly specic to the neuron information transfer function but may be
used as a guiding value for biologically feasible IR electromagnetic
eld intensities. Thus, keeping in mind the total thermal energy in
the volume of the cell 10
20
J and that the detected IR radiation
from nerve cells is 2-3 orders of magnitude higher than that, we
may assume that the energy content inthe cell ininfraredspectrum
of approximately 10
18
10
16
J is within quite feasible range.
6.4. Electric Field Gradients of Biologically Important Modes
Now we know the range of possible energy content in the cell.
Energy of the electric eld E in the volume V can be calculated as
W =
1
2

v

dV (20)
where is relative permittivity, E, and E* is electric eld intensity
and its complex conjugate, V is the volume of the resonator.
We are able to calculate total energy of electric eld of every
mode numerically in COMSOL (based on Eq. (20)). From that we
can deduce the intensity of electric eld E and gradients of electric
eld and vice versa. We have found that to obtain electric eld gra-
dients sufcient to overcome Brownian motion within 100s (using
assumptions in Section 6.2) the energy of the mode has to be at
least on the order of 10
16
J. Gradient of the electric eld squared
(important parameter for dielectrophoretic force) has been calcu-
lated in COMSOL for the energy of mode of 10
16
J and is depicted
in Fig. 7.
In the case of spherical resonator the electric eld of mode of
interest acts by dielectrophoretic force to position centrosome or
other protein rich (high permittivity), large organelles to the centre
of the cell, i.e. the force is centripetal. This location of centrosome
and nucleus is observed exactly in nondividing cell schematically
depicted in Fig. 1. In the case of ellipsoidal resonator, the electric
eld of mode of interest acts by dielectrophoretic force to posi-
tion centrosome or other protein rich (high permittivity), large
organelles to the foci of the cell. This location of centrosomes and
364 M. Cifra / BioSystems 109 (2012) 356366
Fig. 7. (a) Direction (black arrows) and relative magnitude of dielectrophoretic force given by the gradient of squared electric eld

F
2
. White color indicates zones where
E
2
drops below10
16
V
2
m
3
. (a) For spherical dielectric resonator mode (see also Fig. 4, x =0 plane. (b) For prolate ellipsoid dielectric resonator mode Fig. 5, z =0 plane.
nuclei is observed exactly in dividing cell schematically depicted
in Fig. 2. Smaller organelles or macromolecules will be affected
only by stronger elds (gradients) since the dielectrophoretic force
depends on the volume of the particle it acts on, here organelle or
macromolecule.
In both spherical and ellipsoidal dielectric resonator (Fig. 7),
there are zones within the cell volume where the force acts in
centrifugal directions, i.e. away fromthe centre of the nondividing
spherical cell or away fromthe foci of the ellipsoidal dividing cell.
However, the force is rather small in those regions. Therefore, ther-
mal forces or other mechanisms can move the organelle from the
zone of centrifugal dielectrophoretic force tothe zone of centripetal
force.
7. Discussion
The mechanismof large organelle (centrosome and/or nucleus)
positioning by force exerted by electric eld of specic electromag-
netic modes of the cell was explained. However, there are fewmore
issues which could not be covered in current paper and should be
discussed.
First is the issue of minimum required energy in the mode in
order for it to exert sufcient force on the organelle through the
electric eld. Expression for dielectrophoretic force on spherical
particle studiedincurrent paper is givenby Eq. (15). To obtainsuf-
cient force, either the gradient of the squared electric eld (related
to total energy in the cell) or the ClausiusMossotti factor has to
be large enough. For spherical organelle (particle) on which the
dielectrophoretic force is acting, CM factor is limited to the range
<0.5, 1> (Morgan and Green, 1997). If the particle does not have
spherical shape, or its polarization is not isotropic, CMfactor (now
termed polarization factor K()) is not limited by interval <0.5,
1> as for spherical particle but may assume values higher than 1. In
case of highly anisotropic polarization or prolonged shape (such as
prolate ellipsoid with semi-axes a b=c) and long axis of particle or
axis with high polarization parallel to the electric eld, polarization
factor K() reduces to (
p

m
)/
m
)) (Morgan and Green, 1997), i.e.
Eq. (15) becomes
F
DP
=
2ubc
3
Re{
m
K()}
2
(21)
where K() =
p

m
/
m
is polarization factor
If permittivity of the particle (
p
) is much higher than the per-
mittivity of the medium (
m
), gradient of squared electric eld
(E
2
) may be smaller than those described as minimal required at
the end of Section 6.2 while the dielectrophoretic force will be suf-
cient to exert positioning effect on organelles within reasonable
time scale. Large difference (
p

m
) can occur in the case of high
quality (Q1) resonance of particle with external eld. In technical
physics, metamaterials exhibit resonances whichincrease effective
relative permittivity of the material to the order of 10
4
as measured
by Krupka (2008). However, if such analogies of technical metama-
terials exist in biology is just currently being researched, e.g. by
Giakos (2010). If such resonant behavior of organelle or macro-
molecule occurred, this would decrease minimal required energy
content in the electromagnetic mode of interest (also gradient of
squared electric eld) to have biological signicance. According to
Eq. (21), if K() =N, the dielectrophoretic force F
DEP
acting on ellip-
soid is greater by factor N compared to the spherical particle with
same volume and with CMfactor =1, while the gradient of squared
electric eld E
2
is kept same.
Second issue to discuss is the coherence of electromagnetic
modes. In principle, also incoherent eld is capable of excitation of
electromagnetic resonator modes. For the mode of interest in non-
dividing spherical cell, even if the excitation of mode is incoherent
but at the correct frequency, the maximumof intensity of electric
will be always located in the center of the cell and the dielec-
trophoretic force will always act in centripetal direction. Same is
valid for the mode of interest in the case of diving ellipsoidal cell.
Coherence of the excitation would be very important if the input
energy rate (pumping) of the mode was small and the quality factor
of the mode was high so that the energy could sumup. Excitation
should be coherent for efcient summing up of the energy. Other-
wise, if the phase relationships are random(incoherent excitation),
energy would not sumup effectively.
Third issue is the mode selection. In radioengineering, mode is
selected by running the excitation current along the electric eld
line of the mode to be excited and preferentially in the location of
maximumof electric eld of the mode. It may be the case that eld
generators in cell have orientation which prefer the specic mode
dependent on the cell cycle. It also may be that the high excita-
tion of specic mode is triggered only in certain cell cycle phases
or under specic situations to steer the organelle motion. Syner-
gistic mechanisms of these processes, if exist at all, remain to be
discovered. For future theoretical work, the higher modes of the
biological cell dielectric resonator are of the interest. One reason is
that these modes lie in the optical region and it is well documented
that metabolically active cells are source of the optical radiation
(Poppet al., 1992; PoppandBeloussov, 1996, 2003; Beloussov et al.,
2000, 2007; Musumeci et al., 2003; Van Wijk and Shen, 2005; Cifra
M. Cifra / BioSystems 109 (2012) 356366 365
et al., 2011; Rastogi and Pospsil, 2010; Prasad and Posp sil, 2011)
so there is no doubt about the source and excitation of those modes
basedonenergetic reasons. Further, it is well knownthat the higher
modes (WGM Whispering Gallery Modes) in spheroidal dielectric
resonators may reach very high quality factors (Q10
3
10
13
) as
reviewed e.g. by Chiasera et al. (2010) because they are only lim-
ited by dielectric losses and scattering within the resonator. The
high Qfactors of WGM can be easily understood using ray propaga-
tion approach. Higher modes in spheroidal resonators concentrate
their energy near the boundary which corresponds to wave which
is incident on the boundary well beyond the critical angle so the
total internal reection occurs.
9
The WGM reaching the visible
region could realistically have high Q stemming from the WGM
type of propagation and low absorption loss in watery cytosol in
the visible spectral region. They could store the energy and effec-
tively transmit it together with the information along the cellular
membrane. Microtubules can be bound to membrane connecting
the centrosome which is located near the nucleus. Nucleus has the
dimensions to support the dominant electromagnetic modes in the
optical region as studied by (Rattemeyer, 1978; Popp et al., 1989;
Van Wijk and Shen, 2005; Popp and Beloussov, 2003). Mitochon-
dria which are situated along the microtubules have been studied
as optical waveguides by Thar and Khl (2004) and are also source
of photon emission (Batyanov, 1984; Stauff and Ostrowski, 1967).
Electric eld of the specic cellular eigenmodes does not nec-
essarily need to organize the cell organelles only by exerting
the force on the them which results in net translation. Complex
organelles may be sensitive, especially in the case of centrosome,
to the electric eld gradient which could provide the information
about the position. In such a case, organelles must possess mecha-
nisms for the processing of the information while the heavy work
could be exerted by molecular mechanical means by dynamical
cytoskeleton movement which involves motor protein action. This
idea is congruent with the ndings of Albrecht-Buehler, who pro-
posed based on his experiments that motile cells have rudimentary
near infrared vision, where the detectors could be perpendicularly
oriented centrioles in cell centrosomes. Infrared electromagnetic
elds was found to play a role in mutual cellular interactions
(Albrecht-Buehler, 1992, 1991, 2005). For review see (Albrecht-
Buehler, 2010).
8. Conclusion
Eigenmodes of the spherical and ellipsoidal electromagnetic
dielectric resonator have been analysed. The size and shape of the
resonators have been chosen to represent the shape of the inter-
phase and dividing cell. Electromagnetic modes that have shape
exactly suitable for positioning of the sufciently large organelles
in cell (centrosome, nucleus) have been identied.
We analysed directionand magnitude of dielectrophoretic force
exerted on large organelles by electric eld of the modes. We found
that theTM
1m1
modeinspherical resonator acts bycentripetal force
which drags the large organelles with the higher permittivity than
that of the cytosol to the centre of the cell. Specic TM-kind of
mode in the ellipsoidal resonator acts by force on large polarizable
oranelles ina directionthat corresponds tothemovement observed
in the movement of the centrosomes during the cell division, i.e. to
the foci of the ellipsoid cell.
9
This kind of propagation (along the curved boundary with very small losses)
is used to explain the effective propagation of the weak sound (whisper) along the
curved walls of galleries. Hence, they are often termed as Whispering Gallery Modes
(WGM). They were rst observed in acoustic domain. Lord Rayleigh was the rst to
interpret them in terms of efcient reection on the curved walls of the dome of
Saint Pauls Cathedral in London.
Energy supply for the excitation of the eigenmodes has been
theoretically analysed. Minimal required force (10
16
N), gradient
of squared electric eld and corresponding energy (10
16
J) of the
mode that would have biological signicance within the time of
the order magnitude of time required for cell division have been
determined. Minimal required energy of the mode for the biolog-
ical action to take place can be lower in the case of resonance of
organelle with the eld of the cellular resonator mode.
In case of sufcient energy in the biologically relevant mode,
electromagnetic eld of the mode will act as a positioning or
steering mechanism for centrosome and nucelus in the cell, thus
contributes to spatial and dynamical self-organization in biological
systems.
Acknowledgment
Author acknowledges support from Czech Science Agency,
projects n. P102/10/P454 and P102/11/0649. Discussions with Ji r
Pokorn y are deeply appreciated.
Appendix A. Power Losses in Resonator and Related Q
Factors
The power P
d
dissipated in the resonator in the mediumdue to
dielectric losses can be written for both TE and TMmodes as
P
d
= o
d

v

dV = W tan (A.1)
where o
d
is effective conductivity due to dielectric losses and tan
is the loss tangent. RHS of the equation stems fromthe usage of Eq.
(20) and writing o
d
= tan
Resonator quality factor assuming only dielectric losses can be
thus written as
Q
d
=
W
P
d
=
1
tan
=

(A.2)
Appendix B. Mixture of Dielectric Materials
We assume simple model of the mixture of two materials: the
host and the included material (Sihvola, 2000, 2005). Relative per-
mittivity of the more abundant host material is denoted as
e
and
the permittivity of the material included is denoted as
i
. The frac-
tional volume occupied by the included material is f. Then the
fractional volume occupied by the host material is 1f. It is fur-
ther assumed that the particles of included material are spherical
while they need not be of the same size if only all of themare small
compared to the wavelength. Then, the Maxwell Garnett mixing
rule can be used:

cjj
=
c
+3j
c

i

c

i
+2
c
j (
i

c
)
(B.1)
where
eff
is effective permittivity of the mixture of the two mate-
rials.
References
Aas, E., 1996. Refractive index of phytoplankton derived from its metabolite com-
position. J. Plankton Res. 18 (12), 22232249.
Alberts, B., Johnson, A., Lewis, J., Raff, M., Roberts, K., Walter, P., 2008. Molecular
Biology of the Cell, 5th edition. Garland.
Albrecht-Buehler, G., 1991. Surface extensions of 3T3 cells towards distant infrared
light sources. J. Cell Biol. 114 (3), 493502.
Albrecht-Buehler, G., 1992. Rudimentary form of cellular vision. Proc. Natl. Acad.
Sci. U.S.A. 89, 82888829.
Albrecht-Buehler, G., 2005. A long-range attraction between aggregating 3T3 cells
mediated by near-infrared light scattering. J. Cell Biol. 114, 493502.
Albrecht-Buehler, G., March2010. Cell intelligence. http://www.basic.northwestern.
edu/g-buehler/cellint0.htm.
366 M. Cifra / BioSystems 109 (2012) 356366
Arnone, D., Ciesla, C., Pepper, M., et al., 2000. Terahertz imaging comes into view.
Physics World 4, 3540.
Balanis, C.A., 1989. Advanced Engineering Electromagnetics. Wiley.
Barth, A., 2007. Infrared spectroscopy of proteins. Biochim. Biophys. Acta 1767 (9),
10731101.
Batyanov, A.P., 1984. Distant optical interactionof mitochondria. Bull. Exp. Biol. Med.
97 (6), 740742.
Beloussov, L., 2008. Mechanically based generative laws of morphogenesis. Phys.
Biol. 5, 015009.
Beloussov, L., Popp, F., Voeikov, V., Wijk, R.V. (Eds.), 2000. Biophotonics andCoherent
Systems. MoscowUniversity Press.
Beloussov, L.V., Voeikov, V.L., Martynyuk, V.S. (Eds.), 2007. Biophotonics and Coher-
ent Systems in Biology. Springer.
Chiasera, A., Dumeige, Y., Fron, P., Ferrari, M., Jestin, Y., Nunzi Conti, G., Pelli, S.,
Soria, S., Righini, G., 2010. Spherical whispering-gallery-mode microresonators.
Laser Photon. Rev. 4 (3), 457482.
Cifra, M., 2009. Study of electromagnetic oscillations of yeast cells in kHz and GHz
region. Ph.D. Thesis. Czech Technical University in Prague.
Cifra, M., 2010. Biological cell as IR-optical resonator. In: Latin America Optics and
Photonics Conference. Optical Society of America.
Cifra, M., Farhadi, A., Fields, J.Z., 2011. Electromagnetic cellular interactions. Prog.
Biophys. Mol. Biol. 105, 223246.
Cifra, M., Lampa, O., 2008. Simulation of electromagnetic eigenmodes of biological
cavity structures in COMSOL Multiphysics. In: POSTER 2008, Prague.
Cooper, D.E., Powell, D.D., 1986. Infraredproperties of biological materials of interest
to the army. Tech. Rep., U.S. Army Research Ofce.
Cope, F.W., 1973. Electron-phonon (trapped photon) coupling and infrared coaxial
transmission line theory of energy transport in mitochondria and nerve. Bull.
Math. Biol. 354 (56), 627644.
Daga, R., Yonetani, A., Chang, F., 2006. Asymmetric microtubule pushing forces in
nuclear centering. Curr. Biol. 16 (15), 15441550.
Downing, H., Williams, D., 1975. Optical constants of water in the infrared. J. Geo-
phys. Res. 80 (12), 16561661.
Ellison, W., 2007. Permittivity of pure water, at standard atmospheric pressure,
over the frequency range 025THz and the temperature range 0100

C. J. Phys.
Chem. Ref. Data 36, 1.
Euteneuer, U., Schliwa, M., 1992. Mechanismof centrosome positioning during the
wound response in bsc-1 cells. J. Cell Biol. 116 (5), 11571166.
Fraser, A., Frey, A.H., 1968. Electromagnetic emission at micron wavelength from
active nerves. Biophys. J. 8, 731734.
Gastine, M., Courtois, L., Dormann, J., 1967. Electromagnetic resonances of
free dielectric spheres. IEEE Trans. Microwave Theory Tech. 15 (12),
694700.
Giakos, G., 2010. Novel biological metamaterials, nanoscale optical devices, and
polarimetric exploratory data analysis (peda). Int. J. Signal Imaging Syst. Eng.
3 (1), 312.
Hand, D., 1935. The refractivity of protein solutions. J. Biol. Chem. 108 (3), 703707.
Holy, T., Dogterom, M., Yurke, B., Leibler, S., 1997. Assemblyandpositioningof micro-
tubule asters in microfabricated chambers. Proc. Natl. Acad. Sci. U.S.A. 94 (12),
6228.
Hlzel, R., Calander, N., Chiragwandi, Z., Willander, M., Bier, F.F., 2005. Trapping sin-
gle molecules by dielectrophoresis. Phys. Rev. Lett. 95 (September(12)), 128102.
Jelnek, F., Pokorn y, J., 2001. Microtubules in biological cells as circular waveguides
and resonators. Electr. Magnetobiol. 20 (1), 7580.
Julien, A., Guillon, P., 1986. Electromagnetic analysis of spherical dielectric shielded
resonators. IEEE Trans. Microwave Theory Tech. 34, 723729.
Kokkorakis, G., Roumeliotis, J., 1997. Electromagnetic eigenfrequencies in a
spheroidal cavity. J. Electromagn. Waves Appl. 11 (3), 279292.
Krupka, J., 2008. Measurement of the complex permittivity of metal nanoislands and
the surface resistance of thin conducting lms at microwave frequencies. Meas.
Sci. Technol. 19, 065701.
Ku cera, O., Havelka, D., 2012. Mechano-electrical vibrations of microtubuleslink to
subcellular morphology. Biosystems 109, 346355.
Lamprecht, I., 1980. Biological Microcalorimetry. Academic Press, London, Ch.
Growth and metabolismin yeasts, pp. 43112.
Li, L., Kang, X., Leong, M., Wiley, J., 2002. Spheroidal Wave Functions in Electromag-
netic Theory. Wiley Online Library.
Li, L., Li, Z., Leong, M., 2003. Closed-form eigenfrequencies in prolate spheroidal
conducting cavity. IEEE Trans. Microwave Theory Tech. 51 (3), 922927.
McMeekin, T.L., Wilensky, M., Groves, M.L., 1962. Refractive indices of proteins in
relation to amino acid composition and specic volume. Biochem. Biophys. Res.
Commun. 7 (2), 151156.
Mehl, J., 2009. Second-order electromagnetic eigenfrequencies of a triaxial ellipsoid.
Metrologia 46, 554.
Morgan, H., Green, N., 1997. Dielectrophoretic manipulation of rod-shaped viral
particles. J. Electrostat. 42 (3), 279293.
Musumeci, F., Brizhik, L.S., Ho, M.-W. (Eds.), 2003. Energy and Information Transfer
in Biological Systems: How Physics Could Enrich Biological Understanding
Proceedings of the International Workshop. World Scientic Publishing.
Ohki, S., 1968. Dielectric constant and refractive index of lipid bilayers. J. Theor. Biol.
19 (1), 97115.
Pietak, A., 2011a. Endogenous electromagnetic elds in plant leaves: a new
hypothesis for vascular pattern formation. Electromagn. Biol. Med. 30 (2), 93
107.
Pietak, A., 2012. Structural evidence for electromagnetic resonance in plant organ
morphogenesis. Biosystem109, 367380.
Pietak, A.M.,2011b. Electromagnetic resonance in biological form: A role for elds
in morphogenesis. In: Journal of Physics: Conference Series, vol. 329. IOP Pub-
lishing, p. 012012.
Pohl, H.A., 1978. Dielectrophoresis. Cambridge Univ. Press, London.
Popp, F.-A., Beloussov, L.V. (Eds.), 1996. Biophotonics, Non-equilibriumandCoherent
Systems in Biology, Biophysics and Biotechnology. Bioinformservices, Moscow.
Popp, F.-A., Beloussov, L.V. (Eds.), 2003. Integrative Biophysics Biophotonics.
Kluwer Academic Publishers.
Popp, F.-A., Li, K.H., Gu, Q. (Eds.), 1992. Recent Advances in Biophoton research and
its Applications. World Scientic, Singapore, London, NewYork, Hong Kong.
Popp, F.-A., Warnke, U., Knig, H.L., Peschka, W. (Eds.), 1989. Electromagnetic Bio-
Information. Urban & Schwarzenberg, Mnchen-Wien-Baltimore.
Prasad, A., Posp sil, P., 2011. Linoleic acid-inducedultra-weak photonemissionfrom
chlamydomonas reinhardtii as a tool for monitoring of lipid peroxidation in the
cell membranes. PLoS One 6 (7), e22345.
Rastogi, A., Pospsil, P., 2010. Effect of exogenous hydrogen peroxide on biopho-
ton emission from radish root cells. Plant Physiol. Biochem. 48 (23), 117
123.
Rattemeyer, M., 1978. Models for interpretation of photonemission from biologi-
cal systems, in German, Modelle zur Interpretation der Photonenemission aud
biologischen Systeme. Masters Thesis. Universitt Marburg.
Reinsch, S., Gonczy, P., 1998. Mechanisms of nuclear positioning. J. Cell Sci. 111 (16),
2283.
Segelstein, D. J., 1981. Thecomplexrefractiveindexof water. Masters Thesis. Depart-
ment of Physics. University of Missouri-Kansas City.
Sihvola, A., 2000. Mixing rules withcomplex dielectric coefcients. Subsurface Sens-
ing Technol. Appl. 1 (4), 393415.
Sihvola, A., 2005. Electromagnetic Aquametry: Electromagnetic Wave Interaction
with Water and Moist Substances. Springer, Ch. Model Systems for Materials
with High Dielectric Losses in Aquametry, pp. 93112.
Stauff, J., Ostrowski, J., 1967. Chemilumineszenz von Mitochondrien. Z. Naturf. B 22,
734740.
Thar, R., Khl, M., 2004. Propagation of electromagnetic radiation in mito-
chondria? J. Theor. Biol. 230 (2), 261270, http://www.sciencedirect.com/
science/article/B6WMD-4CVV71S-4/2/364319a8a4fa2cb751ff1df1bf7030c3.
Ueda, M., Schliwa, M., Euteneuer, U., 1999. Unusual centrosome cycle in dic-
tyostelium: correlation of dynamic behavior and structural changes. Mol. Biol.
Cell 10 (1), 151160.
Van Wijk, R., Shen, X. (Eds.), 2005. Biophotonics: Optical Science and Engineering
for the 21st Century. Springer.
Vega, J.C.G., 2003. Theory and numerical analysis of the mathieu functions. Tech.
Rep., Technologico de Monterrey, Photonics and Mathematical Optics Group.
http://homepages.mty.itesm.mx/jgutierr/.
Wieliczka, D., Weng, S., Querry, M., 1989. Wedge shaped cell for highly
absorbent liquids: infrared optical constants of water. Appl. Opt. 28 (9), 1714
1719.
Wilmink, G., Ibey, B., Tongue, T., Schulkin, B., Laman, N., Peralta, X., Roth, C., Cerna,
C., Rivest, B., Grundt, J., et al., 2011. Development of a compact terahertz time-
domainspectrometer for the measurement of the optical properties of biological
tissues. J. Biomed. Opt. 16, 047006.
Yadav, R., Singh, I.D., 2004. Normal modes and quality factors of spherical dielectric
resonators. I. Shielded dielectric sphere. Pramana J. Phys. 62, 12551271.