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Articles and Publication  Biology Biotechnologies THE QUANTUM AND CLASSICAL BASIS THE STRUCTURE AND FUNCTIONING BIOLOGICAL MEMBRANES – MOLECULAR MECHANICS OF BIOMEMBRANES
THE QUANTUM AND
CLASSICAL BASIS THE STRUCTURE AND FUNCTIONING BIOLOGICAL MEMBRANES –
MOLECULAR MECHANICS OF BIOMEMBRANES
© S.N. Semenov
Contact: ss.margys@gmail.com
In the beginning was the Word, and
the Word was with God,
and the Word was God.
He was in the beginning with God.
All things came into being through
Him, and apart from Him nothing
came into being that has come into
being.
In Him was life…
(John 1.1 – 1.4)
ABSTRACT.
The paper presented
in popular form the basic concepts of classical and quantum
mechanical foundations of the Molecular Mechanics of Biomembranes
(MMB). Are explained key terms and definitions – the lateral surface
pressure of biological membranes (ΔP),
compressibility (elasticity) of the membranes and phonons – the sound quantum
– quantum mechanical interactions, etc. – used by MMB. It shows what
improvements and/or additions introduced by MMB in a classical mosaic
structure of biomembranes. In particular, it failed to register the any
hydrophobic interactions in biomembranes. But the presence of surface tension
allows membrane proteins to take "hard" conformation, which in the
solution is often simply in principle not been observed. The conditions for the
applicability of the classical and quantum mechanics are considered
to describe different properties of biomembranes. Presented in this article
MMB allow us to consider biological membranes as a mechanical system
obeying to the same time and in one the same place the laws of classical and
quantum mechanics. The boundary between quantum and classical effects is the
velocity occurring in living systems processes. (Bio) phonons that
not only transfer energy from one to another protein but may be while information
signals. It is proposed a mechanism activation and deactivation of various
membrane molecular systems. Briefly describes some of the practical consequences
predicted by MMB and partially confirmed by practice (an action
of alcohol,
local anesthetics,
sound, etc.).
As part of MMB may be explained, and many aspects of the functioning
of the nervous system process, and phonon
mechanism for transmission of nerve
impulses becomes a simple, clear and consistent. This
mechanism includes the appearance of the transmembrane action potential as one
of the intermediate stages of the process. Shows
the potential of MMB for the diagnosis of some cellular pathology and
controlling it,
in particular, to diagnose and to kill cancer cells.
______________________________________________________________________
Introduction
It's no secret that there is now a crisis
in many areas of cell and molecular biology. It’s
accumulated a variety of experimental data, but no, not just
theories, but even hypotheses that allow combining them into a coherent whole.
Without this difficulties arise to the
further development of science, the development of new biologically active
compounds, new methods of diagnosis, treatment, etc. All this is true for
biological membranes. The biomembrane is a critical component of cells – the
basis of all life. Life exists, when living cells exists, when retaining the
integrity of its membrane, both functional and conventional mechanical. It is
known that, despite the variety of cell membranes and their functions, they
consist of two main components: proteins and lipids.
It should be emphasized
that , the
biomembrane is not only an interface between different media, as each monolayer
is. Rather, a biomembrane is an interface separating alive and lifeless. Located
one side of the biomembrane is an alive cell, the other side being lifeless
nature. That is, the biomembrane is located somewhat in between alive and
lifeless. This is a single, known to us, structure with such unique properties.
Despite the diversity of operations
performed, the membranes are characterized by common structural and functional
features. They represent a lipid bilayer matrix in which, on the surface and
which are proteins and other molecules - a common, classical mosaic structure
proposed in the 1935 scientists L. Danielli and G. Dawson ,
which predominated until Singer and Nicolson advanced the fluid
mosaic model in
1972 [1 – 4]. The fluid mosaic model expanded on the Dawson–Danielli
model by including transmembrane proteins, and eliminated the
previously-proposed flanking protein layers that were not well-supported by
experimental evidence.
Molecular Mechanics
of Biomembranes
The main drawback of fluid mosaic model is
that it may not predict new phenomena, not only quantitatively but even
qualitatively. But its main advantage is that it allows its further development,
modernization, which, however, not fundamentally alters the model itself. To
overcome mentioned above disadvantages has been developed, "Molecular Mechanics
of Biomembranes" (MMB). The main achievement
of the proposed MMB is that it allows describing wide range of different
phenomena.
Moreover, MMB
possesses the predictive force. All this was possible to make without the
attraction of any specific assumptions, but only within the framework modern
classical molecular physics and quantum mechanics not only it is qualitative,
but also it is quantitative.
Within the framework classical and quantum
parts MMB naturally are described the influence of lipids on structure
and functioning of the penetrating membrane proteins, the disturbance of
structure and functioning of the biomembranes under the action of outside agents:
antibiotics, venoms, alcohol, etc [6–9]. In particular, the necessity of “fluid
elastic”
lipid phase, the role of lipids in transmission of intermolecular and
trans-membrane information and separation of the latter’s streams, the
mechanism of automatic compensation of internal and external effects and
disturbances become clear.
The aim of this work is the popular description
of the proposed MMB. To do this requires a definition used in the MMB
physical concepts, with their specificities and roles in membrane processes. The
work is intended for those to whom it may help in practical work, especially for
physicians, biologists, chemists, etc. This clarification is necessary because
in this case, some aspects will have to go to the notorious simplification
needed for a popular exposition of MMB.
Presently it is not yet possible to affirm
that the MMB
is completely finished and comprehensive and that it is applicable in all cases.
It is natural that as new experimental facts are accumulated, the MMB
is subject to amendments and clarifications.
This especially relates to “non-classical” quantum phonon
aspects of the MMB, In
particular, issues related to the transfer of information, both in individual
cells and between cells, it does not matter whether it concerns a system
consisting of individual single-celled organisms, or within multicellular
organisms, like our bodies.
Besides, establishment of applicability of the MMB
for viruses containing lipid shells, and possibly to other
lipid-containing viruses needs additional special research. These viruses
comprise many of DNA-containing ones, e.g. pox (variola) and herpes viruses, as
well as all large RNA-containing viruses, except reoviruses. However, the
sufficiently high content of lipids, some 25% of the dry virus weight, which is
comparable to lipid content in plasmatic cell membranes, allows hoping for
successful solution of this issue.
This work is based on previously published
articles devoted to one or other aspects of MMB [5 – 10] and in some
ways is a continuation. In fact, here is a popular introduction to quantum
phonon biology.
Surface pressure
of biomembranes
It is known that lipids are the main
component of biological membranes, forming a lipid bilayer, in which and on
which there are other molecules. The lipids of different cells have different
fatty acid composition and different polar heads. In the beginning restrict
ourselves to the
mane properties of the lipid bilayer.
Previously, I was shown [9] that the
packing density of molecules in the membranes of different cells is different,
which manifests itself in varying value of the lateral surface pressure of
biological membranes ( ΔÐ).
It follows from thermodynamics that
spreading of any surface-active material at the air – water interface lowers
its free energy – this
is the definition of a surfactant. The value of surface
pressure (ΔÐ)
measured in the experiment constitutes the decrement of the free energy (ΔF)
per unit area of the surface. For monolayers –
two-dimensional systems – ΔÐ
and ΔF are equivalent, so we can speak about the pressure ΔÐ
is the pressure as the molecules to each other, similar to the gas pressure in
the tank. Biomembranes can be viewed as a system consisting of 2 monolayers:
internal and external. The presence of different membrane proteins in principle
to the picture is not affected.
The surface pressure is applied to each lipid
monolayer of biomembrane for all molecules in the membrane, as a force tending
to compress and often compresses them. Moreover, such compression is anisotropic
and it acts parallel to the surface of the biomembrane (Fig. 1).
Fig. 1 .
ΔÐ action on membrane
proteins (è )
It is known that state of equilibrium
nonclosed system at constant temperature and volume (area) corresponds to the
minimum of free energy. The interface between biological membranes and washes
its water solution
is an example of such a system. Naturally, this applies to normal conditions,
when the time of observation (measurement) ΔÐ
is smaller than the characteristic time for the existence of cells in a separate
phase of their lifecycle. (For example, it may not relate to the cell membranes
in the cell division process. In
mitosis ΔÐ have to be
changed. Of course, first there was a 1 membrane, and then become 2 membranes.
The process between the existence of a cell membrane and the appearance of the
2nd membrane, by definition, may not be stationary.) Biomembrane surface
pressure ΔÐ is an integral,
average parameter (when the observation time is longer than the time
characteristic of the individual fluctuations in the membrane), which describes
macroproperties the whole system. In such systems, a surfactant displaces (pushes)
from the interface between the membrane-solution less surface-active surfactant.
Schematically shown in Fig. 1, where the dotted red line denotes the surface
activity of a membrane protein that is "Force" tending to remove part
of the molecule from contact with water (polar phase) and convert it into a
non-polar phase – in the region of hydrocarbon chains of lipid molecules.
However, the presence of the surface pressure of the membrane (ΔÐ)
hinders this process.
Within the framework of MMB
it is shown by using the specially developed procedure [9] that the cytoplasmic
membranes of different cells possess the different surface pressure (ΔP),
i.e., with the different density of the packing of molecules in the lipid matrix
of the biomembranes. Cells preserve value ΔP constant with a change
in the ambient conditions, changing the phospholipids composition of its
biomembranes. Values ΔP of a number of the biomembranes are given
below:
- The surface pressure of cellular membranes for
Staphylococcus aureus and Sareina letea
are 55,0 ± 5,0 dynes/cm;
- The surface pressure of cellular membrane for Bac.
subtilis is 20 – 25 dynes/cm;
- The surface pressure of cellular membrane for E.
coli is 20 dynes/cm;
- The surface pressure of cellular membrane for Mycob.
phlei. is 15 – 20 dynes/cm.
- The surface pressure of membranes for Halobacterium
halobium (Bacteriorodopsin’s plaques) is 34.5 ± 0.5 dynes/cm.
- The method is suitable for not only prokaryote
membranes, but also for eukaryotes. The value of lateral surface pressure
for membranes of human erythrocytes is 34,5 ± 0,5 dynes/cm; for rat’s
liver mitochondrial external membranes it is 21,0 ± 0,5 dynes/cm.
To determine ΔP
it is possible to measure or to
control various membrane (cell’s) processes: cell death, change of enzyme
activity of membrane enzymes, ability to perceive the external stimulation of
receptors, etc. The resulting values of ΔP
at the control of various membrane processes gave similar values which
lie within experimental error identical to the membranes of the same cells. This
indicates that the parameters of the lipid bilayer of biological membranes in a
similar way affect at different membrane proteins, i.e. these parameters are
indeed characteristic of the biomembrane own defining its operation.
In such a way non evident link between the MMB
and genetic engineering became visible. So, for example, membrane proteins of
microorganisms Mycob. phlei. may not simply be integrated into the
membranes of microorganisms Bac. subtilis, whereas the reverse situation
membrane protein Bac. subtilis can exist in cytoplasmic membranes Mycob.
phlei. Thus, not every cell is suitable as a recipient in genetic
engineering. It is necessary to take into account the value of the surface
pressure of the cytoplasmic membranes of cells, which are used in genetic
engineering procedures. So, for example, acetylcholine receptors could not be
located in the phospholipid monolayers, surface pressure which exceeds 28 dyne/cm
at pH 7.0, and cholinesterase – in monolayers with ΔP
> 24 dyne/cm [6. Part III, 1 “Quasistatics
Approach”].
In addition, the data presented above make
clear point occurred in hospitals, especially in surgery, "the problem of
staphylococcal infection." Dense cytoplasmic membrane staphylococci,
abnormally dense, so disinfectants do not act on it
– disinfectants penetrate into the membrane can not.
A simple estimate, taking into account the
experimental error in the measurement of the surface pressure membranes of
microorganisms, shows that to achieve the same effect for the staphylococci
aureus (cell death), demands, roughly, from 20 – 50 fold increase in the
concentration of the acting agent, or respectively, the same increase in the
duration time of the agent to a cell as compared to "normal" organisms,
like E. coli. Often, it is simply not feasible conditions – concentrated
hydrogen peroxide just very unstable at high concentrations, surface active
compounds form micelles at high concentrations [11], i.e. when the CMC (critical
micellization constant), which in principle can not achieve the necessary
solution to the concentration (activity) of conventional disinfectants, etc.
The problem requires a radically new
approach to their solution. Analysis of the results and conclusions regarding
the stability of cell membranes and, consequently, the very life of cells
derived from MMB, allowed to find a solution. It is used as a
disinfectant XeF2, solid crystalline compound capable of at
room temperature spontaneously
arbitrarily sublimate, ie, changes in the gas phase. Molecules of this compound
are stable in the solid and gas phases in the absence of water, the presence of
which causes decomposition with formation of a very strong oxidizer F-.
In the dry air and dry surfaces of equipment there is always water in the polar
heads of lipid biomembranes. (Bacterial cells, as well as dust, are the centers
of condensation of gaseous water – a vapor.) This is an area on the surface of
microorganisms, and decomposition of xenon difluoride occurs, i.e.
a local concentration of a strong oxidizer F- exactly where
it's needed, wherever it is required to act by oxidizing components of
biomembranes and causing cell death. Yet been found resistant organisms
surviving after treatment XeF2 [12]. XeF2
may be very useful for situations, when the
agent, which causes infection, is not known and/or there is no time for its
determination. It is easy to sterilize (gas sterilization/disinfection)
different volumes, surfaces and so on by solid xenon difluoride. This
is one the most complicated practical consequence of MMB.
Surface pressure of the cytoplasmic
membrane ( ΔP),
as previously shown, not due to the so-called hydrophobic interactions of lipid
molecules between themselves or with other membrane molecules. Measurements have
shown that if there is such interaction, it is not much, not more than kT
– particles thermal motion energy in the system, where k – Boltzmann
constant and T – absolute temperature. Simply, free energy (F),
where hydrocarbon fatty acid residues of lipid molecules are in contact with
water more than the free energy of the system, where these residues are not in
contact with water. I.e. if you do not like something it does not mean that you
should definitely love something concretes other. Rather, the situation is
reminiscent of the crowd, especially in transport during rush hour: You would
have been glad to take up more space, but others, you may not afford it, even if
they and want to it – there is simply no space available, and necessary to go.
Of course, this is not a scientific analogy, but it’s the simple and obvious.
It should be noted that the term "hydrophobic/hydrophilic"
is meaningful only for sufficiently large systems, when it shall come into force
on statistical regularities, and make no sense when considering the individual
molecules. Correct, this entropy contribution to the free energy of the system,
indicating the probability of the existence of a particular state in a huge
system.
At present there is no way to say, equally
or not ΔÐ
on the inner and outer surfaces of biological membranes. It is likely that the
values of ΔÐ
can be different on different sides of the biological membrane, the more that
they can have a different lipid composition, as, for example, membranes of red
blood cells. In addition, the ΔÐ
and may vary depending on the distance from the surface of the membrane, and the
curvature of the latter. The answer to this question requires further research.
Although brief, this issue will be discussed below. (It should be noted that now
may be measured ΔÐ
only at the outer side of the biomembrane.)
Studies have shown that the forces causing change
in aggregation and/or conformational parameters of membrane-active molecules at
the interface between polar and nonpolar phases (media) are also not some
hydrophobic interaction between them, but a simple mechanical compression of the
molecules – that is, simple mechanical collisions of molecules with each other
in such a system.
Even in densely packed monolayers formed of
mellittin at the interface water-air (at the interface between polar and
nonpolar phases), surface pressure which is comparable to the surface pressure
of cell membranes, the behavior of the molecules obey the Van der Waals
law for an ideal 2-dimensional gas [5, 6, Part 1.2]. Actually,
it was, to some extent, a surprise for me. A dense monolayer, when the molecules
are known to interact with each other, was the ideal two-dimensional gas.
It's already happened as a consequence of that in the
classical hart of MMB biomembranes
easier to consider as a kind of
two-dimensional gas, which lies on a 2-dimensional surface, i.e. surface, which
itself is curved in 3-dimensional space, and thus may have a different curvature
at its various sites. In this case, the change of curvature in the field of
lipid bilayers will certainly change the packing density of lipid molecules,
even more likely not the molecules themselves, but their individual parts.
Obviously, with the convex side of the membrane will increase the area per one
lipid molecule polar head and a concave – to decrease. This is natural, since
hydrocarbon chains of phospholipids also occupy some of the internal volume of
the membrane. It's just simple geometry, which I have tried to depict in Fig. 2.
Fig. 2.
Schematic representation of changes in packing density of individual
moieties of lipid in the bilayer when the bilayer curvature
changes.
(The blue ovals show the area change of curvature of the bilayer.)
That would provide more afford it more
clearly, you can simply place on the table dominoes (you may use cards) as a
variable curvature line – for example, in the form of a wavy line. At the same
chip (card) must necessarily relate to each other (Figure 2.1).
Still, for clarity, one edge of the chips, you can mark – let it be polar
heads of phospholipid molecules. With such a simulation of the curvature of the
surface of the membrane is clearly seen that the change in curvature leads to
the appearance in the membrane of the "free seats", i.e. areas where
the local density changes. It should be noted that the change is not so much the
packing density of molecules in the membrane, but in general, the packing
density of lipid molecules, the individual parts, as in this case. Indirectly,
this will affect the properties of membrane and other molecules. It seems that
this is one of the unique properties of biological membranes.
Reducing the "density" of the
polar heads of lipid molecules leads to a locally reduced in the area of the
surface pressure of the membrane, i.e. by the appearance of a grad Ap. (Where
there's nothing there and the pressure can not be. Nothing in either-something
that does not exist, can not crush.) Therefore, in this area can be accumulated
other, the lipid molecules that form a kind of "islands” (rafts) in lipid
matrix of biomembranes. At the same time, these "islands" can
accumulate molecules surface activity which may be lower than the average value
of the surface-pressure ΔÐ
of the biomembrane, i.e. show areas with specific properties.
Fig. 2.1.
Simulated changes in surface curvature of the membrane by means of chips
( an
explanation in a text).
Thus, begins to play the role of the spatial
correspondence of different membrane molecules of avoiding the emergence of
"voids" in the membrane, which apparently does not improve the barrier
properties of the latter. The necessity for spatial (steric) compliance, the
adjacent membrane molecules observed by other authors [13, 14].
Reducing the "density" of the
polar heads of lipid molecules leads to a local decrease in this area of the
surface pressure of the membrane, i.e. rise to grad
ΔÐ. (Where there's nothing
there and the pressure can not be. Nothing on anything that does not exist can
not press.) Therefore, in this area may accumulate others, not lipid molecules
that form a kind of “rafts” (islands) in the lipid matrix of
biomembranes. At the same time, these "islands" may accumulate
molecules which ΔÐ may be
below the average surface pressure (ΔÐ)
of the biomembrane, i.e. to make areas with specific properties.
The above-described a possible mechanism for the
formation nonlipid “rafts” in biological membranes makes it easy to
identify a number of structural features of molecules that will promote their
"self-organization" in such nonlipid structures at the surface of
various membranes [15 – 19]. A more detailed examination of this issue may
give a simple mechanical explanation of the possible processes change the shape
of cells, possible mechanism of their movement. The last issues are especially
characteristic of microorganisms and protozoa. For example, to describe the
movement of cells in the direction of grad nutrients. In other words, may
not be nor any specific interactions between molecules in these rafts –
there simply is "free" place for these molecules. Here they are, and
it filled. But in present paper I will no longer dwell on them, and some other
particular cases MMB.
It have been already noted that even
densely packed monolayers of membrane-active compounds, in particular, mellittin
at the interface between polar and nonpolar phases are described by Van der
Waals law
for two-dimensional ideal gas, which has an area occupied by a molecule on the
surface. A characteristic feature is their compressibility of gases. By
definition, Compressibility (bulk modulus) of a substance reversibly
changes its volume under uniform outer pressure. Characterized by the compressibility
β, defined as
the relative change in volume V (or density ρ) of
substance with a change in pressure P:
Compressibility of gases, being very large
at pressures up to 1 kbar, as they approach the density to the density of
liquids is close to the compressibility of liquids. The latter, with increasing
P decreases sharply at first, and then changed very little. The linear
compressibility of anisotropic materials depends on the crystallo-graphic
directions (at least up to pressures of tens of kbar), and the directions of
weak interatomic interaction, it can be 8 – 10 times greater than the
compressibility in the directions along which the crystal lattice has a stronger
relationship. Compressibility – the most important characteristic of a
substance that gives an indication of the dependence of physical properties of
the interatomic (intermolecular) distance. The compressibility is closely
related to elastic (elasticity) – a property substance to have the
mechanical resistance to external force and take after it fall the original form;
that is especially true for gas. The
opposition to
elasticity is
called plasticity
[20].
The above definition of the compressibility
for three-dimensional structures easily transforms and for two-dimensional
systems witch are the biological membranes. Sufficient to replace V –
volume by the S – surface area of the membrane, and, for simplicity, that
would not change the current in monolayer measurements, notations remain for the
surface pressure ΔÐ
its designation, the magnitude of the changes of the surface pressure is denoted
as ∂∆Ð
, we get:
We need only this type of formula to
describe the compressibility. In addition, for the lipid may
it be used in the formula, instead of S – the area of
the membrane surface of the value of S' – the area per one molecule on
the surface. Than, membrane area S = nS', where n – number of lipid
molecules on the surface, respectively ΔS
= nΔS'. After making these substitutions, we obtain the same formula,
but at the molecular level. In the future we will use the formula for the
compressibility, but not longer considering the entire area of the biomembrane,
and the only one molecule area in the membrane. Neither base definitions, nor
the numerical values not change, but it is more in line with those phenomena
that we consider in the membrane, because we consider them at the molecular
level.
Need always remember that the biomembrane is
anisotropic system and its compressibility depends from the directions.
Lateral compressibility is very different from the "vertical
compressibility", i.e., the compressibility, the pressure (usually
hydrostatic) acts perpendicular to the surface of each of the two monolayers
that form the membrane. In this case the hydrostatic pressure is much less
effect on its properties of biomembranes [6. “Quasistatics Approach”; 21 –
23]. These biomembranes are somewhat similar to solid crystalline body, which,
as noted above, the compressibility depends on the direction. Bilayers, though
very thin, but 40 – 60 Ǻ possess. Now this is only an analogy, not
playing a significant role in the considering molecular and mechanical
properties of biological membranes in terms of classical physics. Membranes
contain different lipids. It was found that under normal conditions, the
majority of membrane lipids in liquid-crystalline state. (Here I use the
currently accepted terminology.) For example, under normal conditions, the
growth of microorganism’s membrane Micoplasma laidlawii, Micrococcus
lysodeikticus and Escherichia coli form a "liquid
phase", characterized by high mobility of the hydrocarbon chains of lipid
molecules. In this case there is strangeness: temperature decrease of the medium
leads the appearance the crystalline properties lipids, but the same properties
occur for the lipid molecules in the monolayers during compression at the air\water
interface, i.e. when its temperature increase – a paradox. To solve this
problem back to the situation discussed above.
The above data were given to the values
of ΔÐ
for different cells. For most cells, these values are
characteristic for lipids or lipid mixtures, when they have a high
compressibility. To do this lipids are needed, containing hydrocarbon chains of
at least one C = C bond. In addition, the compressibility of the
membranes is necessary to ensure the ability to change their curvature (see fig.
2 and 2.1). But the high compressibility this is the gas property, not fluid. In
this case, the above mentioned paradox disappears. Increasing external pressure,
temperature reduction are processes long used in the liquefaction of gases.
Therefore, we need to say that biological membranes contain the gas and
condensed lipid phases, which corresponds to the experimental results
obtained previously [5, 6, 9].
Temperature change alters the fatty acid
composition of phospholipid molecules that make up cell membranes. An increase
in temperature leads to a decrease in the proportion of unsaturated fatty acids
and decrease – to increase their share. Similar changes in the phospholipid
composition of biological membranes, except for temperature, and can cause
introduction of the hydrophobic region of molecules, breaking microviscosity
biomembranes [24 - 26]. This is an indication that the living cells tries to
maintain a constant surface pressure of its membrane as conditions change.
Increase in the proportion of unsaturated fatty acids decreases membrane
microviscosity [27], i.e. keeps the membrane lipids in the "liquid
phase – traditional terminology or gas phase – my interpretation”.
In this case, the maximum effect is achieved by replacing saturated fatty acids
on the fatty acid with one double bond. A further increase in the number of
double bonds leads to a slight increase in membrane fluidity [22].
A note
associated with the definition of various parameters: the viscosity
(as well as microviscosity) is introduced to describe the motion of the
body (like microbodies) in the liquid isotropic physical environment, which is
not a membrane. It has been noted above. Biomembranes – singular, anisotropic
system in which at a distance, or even less, one of its molecules, possible to
see properties of various physical systems. These different physical systems may
be more in several states of aggregation. This will be discussed below, when
considering the quantum phonon aspects of the membranes.
The lateral surface pressure reflects the packing
density of molecules in the membrane. The lower lateral surface pressure, ie the
lower the packing density of molecules, so, naturally, lower microviscosity of
the medium. Because of a decrease in the packing density of molecules in the
system, we are approaching the void, and emptiness is nothing to resist the
movement of molecules, probes, which are used to measure the viscosity of
various membranes. With this in mind and we may say that the surface pressure
and the "microviscosity" of the membrane close to each other. This
closeness is correlation between them – the parameters are changed similarly.
Summing up we get the view expressed that the
presence of surface tension allows membrane proteins to take "hard"
conformation, which in the solution is often simply in principle not been
observed.
In addition, roughly one may say that,
since the pressure force acts on any particle in the membrane in a direction
perpendicular to the surface of the membrane molecules, then for all the protein
molecules to form a hydrophobic region, which is embedded in the membrane
differs from that of a constant area "lateral cross-section "(Fig. 3),
appear buoyant force (Fc). This force tends to remove
particles (molecules) from the interface mebrane
– in a water environment. And the force of ejection may be directed to the
center of the membrane (not shown). It is this process and counteracts the
surface activity of the membrane protein. In addition, unstable molecules with
an area of "lateral section" will disrupt the structure of lipids, but
around him. This is amply illustrated in Fig. 4, shown on the website of the
Institute of Mathematical Problems of Biology [27].
Fig. 3.
The action of the lateral surface pressure of the membrane ( ΔÐ)
on membrane proteins. (Explanation in a text).
In other words, for the lipid will arise a
situation similar to that discussed in Figures 2 and 1.1, which may lead to a
local change in curvature of the membrane and/or the formation of "islands",
etc.
As already noted, the presence of the
lateral surface pressure acting on the protein molecule in the membrane, as well
as any other molecule that leads to the appearance of the last new "hard"
conformation, which is characterized by the additional the surface
conformational energy. Additional energy is equal to the work on the compression
of the membrane component of the zero surface pressure to a pressure equal to
the pressure of the surface of the membrane. We consider a situation that is
already discussed in connection with the phases of the cell ΔÐ
life cycle. In these circumstances, we may consider a system at equilibrium and
stationary. Then the additional energy derived molecule in the membrane due to
its compression from the environment (El) is:
Fig. 4. The
influence of proteins on the structure of the surrounding lipid bilayer.
The limits of integration from 0 to Pm,
Where: P (S)
– a function of surface pressure of monolayer area S per single
molecule in a monolayer;
Pm
– the equilibrium surface pressure of the concrete biomembrane;
θ
– coefficient taking into account the geometry of the molecule (roughly –
its perimeter and/or contact area of the molecule with its surroundings inside
the membrane) and used the physical system of units.
In practice, to assess the additional
conformational energy can be used so that it equals the area of the
figure bounded by the compression isotherm of a monolayer of fully extended
state to a surface pressure equal to the surface pressure of the biological
membrane ( Ðì),
line P = Ðì
and the y-axis (Fig. 5). That is, additional energy is equal to the work that
must be expended, that would compress the monolayer of a rarefied state where
almost ΔÐ practically
= 0, to a pressure corresponding to the
surface pressure of the biomembrane. It is obvious that in the stationary case,
we are considering the pressure, which the membrane presses on its molecule,
identical pressure, which pushes the molecule to its surroundings. Further, it
is easy to show that you may use to assess the compression isotherms of lipid
monolayers, which is technically much easier to do than to receive and compress
the monolayers of other membrane molecules. In addition, for nonlipid membrane
molecules (proteins, mostly) of their additional energy (Ånl)*
will be equal to the experimentally obtained additional energy
one lipid molecule (Ål)
multiplied by the ratio of areas of the surface membrane in its natural, natural
ΔÐ nonlipid molecule (protein),
(Snl) and average
area occupied by phospholipid molecules (Sl)
at ΔÐ, equal to the normal,
natural pressure of the biomembrane ΔÐì,
line Ð = Ðì
and the y-axis (Fig. 5).
(* Å nl
= Åë Snl/ Sl)
Fig. 5.
For explanation, refer to the text
This figure 5 was probably even more
physical meaning than it might seem at first sight. Really it depicts gas,
rather condensed, but the gas. This is gas, as in monolayers of mellittin. For
them, it was showed, even when ΔÐ
equal to or more surface pressures of cytoplasmic membranes of living cells the
mellittin behavior is described by valid for ideal gases.
Moreover, from the monolayer experiments,
we have that the increase in the degree of unsaturation fatty acid chains of
molecules of phospholipids, other conditions being equal, leads to an increase
in additional energy compression for particles in biological membranes
containing these lipids (increases area of the shaded figure in
Fig. 5). Consequently, the increase in the proportion of unsaturation of the
hydrocarbon chains of lipid molecules leads to an increase in additional
conformational energy of membrane protein for a given surface pressure of the
biomembrane. That is, possible increase the energy of compression of the
membrane, with whom it acts at the penetrating membrane proteins, without
increasing the lateral surface pressure of the membrane
itself.
In other words, the presence of
polyunsaturated fatty acids in biomembranes produces gains in the value added to
the conformational energy without increasing the surface tension of the membrane.
Thus, a compromise may be reached between the relatively low surface pressure,
membrane proteins, allowing the latter to penetrate, embedded in the membrane at
the interface and be there for a long time needed for cell activity, and an
additional value of the conformational energy of the protein molecule, providing
it’s optimal structure.
Different experimental data confirm this
conclusion (see comments above about the correlation between microviscosity and
surface pressure of biological membranes), membranes of cells are characterized
not only certain surface pressure, and varying compressibility, i.e. dependence
of various changes in membrane surface pressure as a function of the packing
density of molecules in it.
By the way ,
this implies that the lipid matrix of well-functioning of biological
membranes may not be "crystal". Because of under the "crystallization"
membrane lipids decrease sharply the value of additional compression energy
obtained by penetrating membrane proteins. In this case the additional energy is
several kT per molecule in the membrane and is comparable with the energy
of thermal motion (k – Boltzmann constant, T – absolute
temperature in degrees Kelvin). Therefore, if the membrane under normal
conditions will contain lipids in the "crystalline" state, the
temperature fluctuations (in other words - the thermal noise) will cause
fluctuations in the structure of membrane proteins – their dramatic change in
the state – and therefore, abrupt random changes in their activity. These
random changes in the activity of membrane systems, of course, the phenomenon is
incompatible with the very existence of life. You can say simply, in the normal
membrane lipid phase transition in the "crystalline" state leads to a
membrane protein penetrating the possibility of "ravel
out or to get an inactive state",
i.e. change their structural state and inactivated.
It is just to remind once again that the presence
of double bonds in the hydrocarbon chains of lipid molecules leads to an
increase in the lateral compressibility of the biomembrane. More precisely, it
is true for the one half of the membrane, where these relationships are present.
Under this model, as will be clear later, this statement is correct, ie membrane
may be considered as a system consisting of lipid matrix and are in it of
membrane proteins. In turn, the lipid matrix itself is also a complex system
consisting of two subsystems – the lipid monolayers.
Therefore, the answer to Prof.
D. Chapman, FRS question: “What are the double bonds required for?”
should sound somewhat like this: “They provide for existence and support of
lateral compressibility (elasticity) of biological membrane, which is
responsible for surplus lateral compression energy received by penetrating
membrane proteins from the biological membrane lipid matrix surrounding them, and
damp the random variation of the lipid molecules density in biomembranes”.
From summarized in this paper, based on
classical physics, part of the MMB implies that the incorporation of
foreign molecules (structural defects) should lead to a change in packing
density of molecules in the membrane, i.e. to a change in its surface pressure
and (lateral) compressibility. Therefore, in the early stages, when there is no
lysis of cells must vary activity of membrane proteins, as evidenced by both
their own experimental results and literature data. More details are described
earlier
[5. “Êâàçèñòàòèêà”;
6. “Quasistatics Approach”; 29 – 33].
In these papers presents data on the effect
of changing the structure of the lipid in the photochemical cycle of
bacteriorhodopsin in the fragments of photosynthetic membranes of halophilic
microorganisms Halobacterium halobium, gas, sound, alcohol, local
anesthetics and other organic solvents, at the cell membranes
[29 – 33]. After the publication this article with the
consideration an alternative mechanism of action of alcohol on cellular
membranes published an experimental date supporting
[34] my conclusion that the abrupt discontinuation of alcohol may be dangerous
to health.
Also, do not want to be right with the
prediction obtained as a result of consideration of sound
action on living organisms. It was found that the observed
number of road accidents, with no apparent reason, may be associated with a
specific narcotic effect of loud music, loud drums containing sounds
[33]. However, this phenomenon has not been studied specifically.
It should be noted the potential danger of
fetal ultrasound
investigations of pregnant women. Fetus is booming, and the
ultrasound frequency range, used for research, not specifically controlled. (This
will be discussed below.) Therefore, in principle, it may effect on cellular
processes of the fetus. However, this requires special studies.
Based on the results within the framework of MMB
possible make some practical recommendations in addition to those already
described in this paper. For example, in order to reduce the possibility that
the "dissolution" of gases in the cell membranes during underwater
work or sport may be recommended diet high in vegetable oils, etc.
It should be noted that even under this approach
is understandable, another biological role of polyunsaturated fatty acid
residual in phospholipid molecules in biological membranes. They are due to the
weak dependence of the surface pressure from the area per 1 molecule, i.e. high
compressibility may smooth out fluctuations, changes in surface pressure
fluctuations in biological membranes, i.e. enhance the stability of various.
Thus, the high compressibility dampening of
the membrane surface pressure pulsation and thus increases the stability of
conditions in which the functioning of various membrane living systems – membrane
molecular systems. By the way, at a
constant temperature of a living system to do it easier. And it is not so
important that a high or low temperature – most importantly, what would it be
stable. And it is not so important that
a high or low temperature – most importantly, what would it be stable. This
could be achieved, or maintaining a constant temperature of his body, becoming a
warm-blooded that is inaccessible the simplest, or living with a constant
ambient temperature. By the way, the rapid development of plankton observed in
the polar regions of the ocean, where there is constant water temperature ~ 4°
C.
Summarize all the above, we have that
biomembranes are highly dynamic structures, because their constituent molecules
are in constant motion. However, despite this flexibility, the lipid portion of
the membrane itself is an excellent insulator, and just a great barrier to free
penetration through it for various molecules. Moreover, such a delicate and
dynamic barrier, visible only through an electron microscope, has a very complex
internal structure, largely obeying a simple law of thermodynamics and classical
mechanics, describing the behavior of large multicomponent system. Of course, we
must understand that this is said with some degree of simplification, made
to facilitate understanding of the material.
To describe the properties of biomembranes
as systems that contain a huge number of different molecules in the framework of
classical MMB, be aware that they have their own value of the surface
pressure ΔÐ.
This value is different for different cell membranes and for membranes of
cellular organelles.
Biomembrane surface pressure allows be in it only
to the membrane molecules (proteins, etc.), surface activity (more precisely,
the maximum surface pressure) is not lower than the surface pressure of the
biological membrane. This applies to the proper membrane components, as well as
to outside molecules.
On each molecule within the membrane of the
lateral compression of the acts a force from its environment tends to compress
and compressed the molecule. This force is similar to that which occurs during
the mechanical collision of molecules at the interface between polar and
nonpolar phases – in a "two-dimensional gas". As a result, the
molecule is having some intramolecular "stress" and/or restrictions,
which are absent in a homogeneous environment – i.e. in the solution.
Such an analysis is possible when we look
at the big multi-component system without taking into account local fluctuations
and/or changes in the membrane related to the operation of its individual
molecular systems. I.e. when we consider the biological membranes as a
stationary system, and therefore was previously introduced the term " Quasistatics"
to emphasize that we nevertheless consider the dynamic structure seems to be
static and equilibrium. Usually it is the processes occurring in a relatively
long period of time > > 10-4 – 10-3
sec.
Quantum-phonon approximation
Let us examine now a different situation
when we study the behavior of individual molecules and may
no longer neglect occurring therein changes. But at the same time, we may ignore
similar changes in the surrounding molecules. I.e., we consider the local
changes already in the membrane, changes in a single molecule or molecular
ensemble in the operating process. Such a division is artificial, since all
processes in the membranes occur simultaneously, and, of course, has its limits
of application.
The need to consider processes occurring at
the level of individual molecules is obvious. And the need to consider processes
on the mechanical level, due to the fact that an increasing number of detected mechanodependent
processes in biological membranes, including ion channels and
operation of excitable nerve cells [35 - 41]. It is not necessary to explain the
importance of understanding the processes of information transmission and it
processing for understanding the basics of life of any organism.
These issues are considered in non-classical,
quantum-mechanical part of the MMB. Indeed, many effects around us look
like classical because classical physical laws are actually based on quantum
mechanics [42]. The need to consider quantum effects arise in recent years in
addressing various aspects of molecular biology, the genome [43], brain and
other events listed, such as “Wikipedia” [44]. Quantum phenomena let be
quantum phenomena, but I’ll use here classical mechanics, without it does not
work. Schrodinger equations are omitted, but since there is such terminology, we
shall, so far, to use it. Moreover, calling something by quantum mechanics (that
we do not understand, can not imagine, call magic, or as here, the quantum
mechanics), we find that the MMB, predicts that the intermolecular
interactions in the membrane must be accompanied by the emission or absorption
quantum “mechanical interaction” – phonons – quantum quasiparticles.
First need to understand what is a phonon. This
term originated in solid state physics. Very good, but not quite clearly its
definition is given in Wikipedia, and the in other works related to solid-state
physics [45 – 47]. At first glance it might seem that the solid crystal has no
relation to biological membranes. But again I must reiterate that
biomembranes – a unique system – separating the living from the nonliving,
other systems with similar properties are not known.
Summing up the various definitions, anybody
can write a phonon – a quasi-particle, representing the quantum of the elastic
fluctuations (vibrations) of a medium. The concept of phonon plays an important
role in describing the properties of solids: crystals lattice the thermal
properties are similar to the gas of phonons [Great Russian Encyclopedic
Dictionary]. Vibrations of atoms in the crystal are replaced by the spread in
the medium of sound waves and quantum which are the phonons, but this is true
for relatively high-intensity sound. And what kind of wave may say, if it may
not be, but there was just variations of one or only a few atoms. Fluctuations
of a single atom in the crystal lattice necessary to some extent, affected at
the neighboring atoms. Phonon has an energy and a mechanical impulse. The
propagation velocity of phonons in the system is equal to the speed of sound in
it. Phonon spin is zero. Phonon belongs to the bosons, and is described by
Bose-Einstein statistics. Unlike ordinary particles, which exist on their own,
including the empty space, quasiparticles can not exist outside of the medium
fluctuations that (or just parts of which) they are. Thus, the phonons are
always elements of the system, no
biomembranes – no phonons.
As well as ordinary particles,
quasiparticles may be more or less localized in space and maintain this
localization in the motion, i.e. have a quasiform, but possible call it a
quasisize or quasivolume. It is really important problem, but in this work it
just indicated. But, more importantly, that the phonon, whatever it may be, has
the energy and mechanical impulse – simply put, it may hit a hurdle and give
him its energy and its mechanical impulse. Let
me explain
it in
simple examples.
Again turn to the dominoes that are set on their narrow ends
well (mentally or actually), as is done during the competitions, where there is
a problem to install as many pieces of dominoes, and then, pushing one, to get
as more interesting drop all the rest.
Fig. 6 shows just such a situation, however, a
much smaller scale.
Fig. 6.
Falling pieces of dominoes.
Somewhere in the turquoise-blue
circle is a phonon itself.
The arrow shows the
direction of the phonon motion –
transfer energy
and its mechanical impulse.
Dominoes, falling, are becoming mechanical
impulse, which then, in a collision, pass the chips worth, and they, in turn,
falling, pass it on.
So the momentum (mechanical impulse) is moved along the system,
passing the initial push, kick, along the whole line from one end to another.
Incidentally, in this system, the transfer of momentum is
practically without losses (if you put all your chips neatly enough), because
each chip has its standing potential energy, because we do it well placed. In
the fall of the potential energy turns into kinetic energy of falling chip. This
process is repeated until as long as there are chips. But it should be noted
that this is a one-time system. All the chips will fall, and the more the system
will not work.
Not difficult to make a system suitable for
multiple operations. Enough to attach to the floor (bottom) chip on a spring or
other elastic fastening. Then the chip after a fairly strong shock will push the
next and back to its original position, and the next push and return the
following to its original position, and that the following ....
This is an ideal situation, but we live in the real world, so
the part of the energy transferred from the chip to chip, with the impact goes
into heating the springs and so go into heat and lost. The energy will be lost
and the impact force transmitted to each other chips to be smaller. A little
earlier or later, the momentum transfer process will fail. What would be the
signal transmitted, the system must have amplifiers along the line of chips.
These amplifiers, having a weak push from one chip should give a strong signal (push
kick) to the next. Naturally, this process will require the expenditure of
energy to amplifier. Understanding these simple processes we shall need later on,
during the transition from Domino to the atoms of membrane molecules in the
transition from classical mechanics to quantum mechanics of phonons. But there
is nothing complicated in this case will simply replace the dominoes on the
balls of atoms, interconnected by elastic springs, as shown in Fig. 7.
Fig. 7
Schematic phonon motion in the crystal structure –
b alls
connected by dark blue and gray lines, and in lipid
– balls
(CH2 group), combined with dark blue lines (gray
lines are ignored.)
The red arrow ()
– the direction of energy
transfer and phonon momentum –
motion of a phonon. Red oval
–
phonon localization in the membrane (the crystal).
In this figure (Fig. 7) attempted to
portray the spread of the phonons in the regular structures. Schematically
considered the crystal structure, where atoms are depicted as circles, connected
by links, represented by dark blue and gray lines, so that the cells form a (two
dimensional) crystal lattice. In a normal crystal lattice, the atoms in the
lattice nodes strongly associated with their neighbors. At the same time as, in
biological membranes, from the viewpoint of thermodynamics and molecular physics,
we can find areas with a "crystalline" structure, located by the polar
heads of phospholipids, and spectacular 7 - 9 C-atoms of the hydrocarbon chains
[5, 6, 48], characteristic of solid crystalline materials at the same time, the
central part of the membrane, as the authors note, has a "disordered"
state, close on the properties of fluids, or maybe even gases. Of course, should
be understood that this is said with some degree of simplification, made
to facilitate understanding of the material. In this paper, I will
not focus on the structural and phase characteristics of individual sites,
regions of biomembranes.
Of course, here will use normal physical terms with the prefix
"quasi" or otherwise "explaining" terms. Something
likes this – a two-dimensional quasigas
on three dimensional surfaces with variable curvature. In this quasigas
may only consist of real particles – molecules, quasiparticles – phonons and
possibly from their mixture. However, consider these questions – the task of
subsequent studies, developing and clarifying the MMB. And now it’s
fundamentally not affecting the essence of the material presented, nor for
deriving conclusions from it.
Vibrations of individual atoms in a regular
structure, such as a normal 3-dimensional crystal, or in the membrane
quasicrystal necessarily capture their neighboring atoms, i.e. process is the
dissipation of energy that goes into the usual thermal energy, i.e. is precisely
to fluctuations neighbors. Thus, in his moving biofonon in biomembranes
has been losing some of its energy, which means that the wavelength of the quasi
particles increases as in quantum mechanics:
Ef = ħ ω
Where – Ef – phonon energy;
ω
– own quantum phonon frequency is inversely proportional to its "wave
length";
ħ
– Planck’s constant.
Therefore, if somewhere in an organism it is
necessary to transmit the information signal in the form of a phonon, it will
have to periodically increase and correct, otherwise it is a final point, if it
can reach it, – where the recipient is to be very different from signal, which
is sent by the sender. This is one of the foundations of the theory of
information transfer. This conclusion is we need in the future when considering
the mechanism of information transfer in living systems.
At this moment, it is popularly presented all the
necessary information on the quantum mechanics of phonons, which is needed in
the future.
MMB – phonons
It was mentioned above that reveals more
and more mechanodependent
phenomena in biological membranes. Under described MMB is a natural
phenomenon. It has already been shown that the biological membrane is subject to
certain laws of classical mechanics and thermodynamics, which, in turn, are
based on the same laws of classical mechanics adapted to systems containing a
huge number of particles.
Based on the above stated it is clear that
incorporation of foreign particles in biological membranes should lead to their
"breaking up" due to the formation of structural defects of lipid
bilayers. At the same time defects are changing its lateral pressure and
compressibility. Therefore, in the early stages, when there is no lysis of cells
must vary activity of membrane proteins, which is confirmed by published data
obtained for gramicidin S and its analogs are used to determine the surface
pressure ΔÐ
various biological membranes [5, 6]. In this case, the most complete effect of
the change ΔÐ was obtained
for Bacteriorhodopsin, which shows that the ability of the protein returns to
its initial state after it is activated by light, decreased with the appearance
of structural defects in lipid plaque. Consequently, the driving force behind
the relaxation of the protein opsin (the main protein Bacteriorhodopsin) in the
initial state has been pressure on it by its membrane environment. In other
words, the excitation of the membrane molecules leads to
the increasing its area in
the membrane. Below, this process will be discussed in more detail as an example.
Further, it has been confirmed in other objects
belonging to different biological membranes. As an example, we note that the
molecule of the Ca-ATPase sarcoplasmic reticulum may be in the 2 conformational
states [49]. In the process of protein when it is activated protein distances
within the active site increased 10 – 20 Ǻ. And then, when you
return to the original state of the protein, the active center decreases again
[50 – 53]. This indicates that the proposed molecular mechanism of mechanical
activation and inactivation of membrane proteins is universal.
It should be noted that the conformational
transformations in the molecules, especially those associated with vibrational
excitation, occur very quickly. The time required for this is the order of 10-8
– 10-12 sec [53 – 55].
These facts suggest that the process of
functioning of the protein in living systems is accompanied by absorption or
emission of quanta of mechanical interaction – phonons. I.e. during the
activation of membrane proteins under external influences such as the binding of
the membrane enzyme with a substrate from
the surrounding solution, and the formation of substrate-enzyme complex energy
is released. Which then, upon return of the protein molecule and the initial (unexcited)
state can be emitted in the form of membrane-phonon or otherwise, for example,
in the form of a photon – bio/chemluminescence. Liberated phonon may then
activate another protein in the membrane system, being absorbed by it, which,
then, having finished their molecular cycle, may also provide a phonon, etc. I.e.
it is a process transmission of information along the membrane and coordination
among the various membrane protein systems. And it is not necessary that these
membrane systems have been spatially close to each other or had any other "visible"
link. May not be apparent coupling of different membrane protein systems among
themselves. We'll just watch an event, usually caused by some effect (or effects),
including a different nature, in a remote section of the biomembrane. The main
thing that existed in the membrane an ordered structure, similar to the crystal
structure, providing conditions for the transfer of phonon information, such
structures, as noted above, exist in biological membranes. In some ways it may
resemble a Christmas tree garland, when it consistently (or alternately, that
does not matter) various light bulbs are lit.
Naturally, the reverse process is possible: the
membrane molecule (in the formula designated "receptor"), absorbing a
phonon, passes into activated state, and then, under the influence of lateral
pressure of the biomembrane is returned to its original, inactive state (or
conformation). These processes may be schematically represented by the following
schema, which describes the reactions that go with the absorption or emission of
energy:
Direct reaction is
(external chemical signal) + (membrane
receptor)è
(a complex molecule with a
receptor signal) + (membrane phonon = ħω);
The reverse reaction:
(a complex molecule with a
receptor signal) + (membrane phonon = ħω) è
(membrane receptor) + (a chemical
signal, extracted from the membrane).
And outside, as well as an internal signal may
not necessarily be the chemical signal. This may be a quantum (s) familiar to us
light, and another phonon (s), temperature, mechanical impact, which is true at
the level of the membrane will not differ from the temperature and
electromagnetic field, etc. (various, including other outsiders, impact, marked
"?", but which may not be). It is more correct to write the
above equations are summarized as follows:
(external signal) + (membrane
receptor) +?
(a complex receptor with a signal)
+ Σ
ħωi
+ Σ hνj
+ ?
Where ħωi
– phonons; i = 0, 1, 2 ...
hνj – photons; j
= 0, 1, 2 ...
ħ (h) – Planck's constant (different
types of notation)
Σ – a mathematical
summation sign.
However, for this popular exposition of the
differences between the given formulas, and formulas themselves are not
important. Simply, they will encounter in other studies, including those of MMB,
is therefore appropriate to explain their meaning here. In the second equation
should be emphasized that there may not necessarily released phonons, but the
light photons, just the energy for heating the body, certain
chemical reactions take place, all together, and certain combinations. In other
words, perhaps the mutual transformation of different signals to each other: the
chemical reaction may cause the phonons and/or photons, and vice versa. I.e., as
in solids, in biological membranes may interconversion of "light" in
"sound" – the effect of sound luminescence.
At
chemical reactions do not stop, about them and so much has been written.
Photons:
there is no reason to prohibit the excess (or specific) energy in the
some membrane process of in the form of light – or rather in
the form of photons. Bioluminescence, which was opened in the first third of the
twentieth century. [56, 57], already is a known and studied phenomenon. This
effect was observed for cells of different nature, as for the cells of
prokaryotes and eukaryotes, animal and plant cells [58 – 63].
Put forward various hypotheses about the role
of biophotons into cellular activity. According to one hypothesis biophotons
remove excess energy that arises in the process of cell activity. According to
other hypotheses biophotons participate in intercellular exchange of information.
Both hypotheses do not contradict the proposed here MMB. But now I will
not stop there. It's just another little explanation showing MMB
relationship with other areas of cell biology, biochemistry, and different
sections of physics, including biophysics, etc.
Conclusions about the role of phonons in membrane
processes have been confirmed experimentally. For this purpose, special
phonon spectrometer designed to obtain the phonon spectra for a number of
cells, which are given in previously published papers [7 – 8]. Fig. 8.
Fig. 8.
Part of the ultrasonic spectrum cells Bac. subtilis in aqueous suspension
(physiological solution)
Along the axis A of the spectrum is given in
relative units cu
Ff – natural frequency of the phonon absorption.
Let us consider in more detail the
formation of phonons in biological membranes with the structural changes in
penetrating the membrane proteins. It is easy to show based on the geometric
parameters of protein molecules, that even "minor" structural changes
in them may affect on the border of the protein-lipid hydrocarbon chains of
membrane area at the border with more than one CH2-group of the lipid.
This is easily done, for example, action with the use of
spatial models of atoms. Already noted above that
for the Ca-ATPase molecules in the linear dimensions of the process of
activation – deactivation alter at 10-20 Ǻ [49 – 52].
Therefore, the protein molecule in the process of its
operation could "push" just a few of the CH2 groups in the
hydrocarbon chain as a single lipid molecule or several at once. I.e. will be
formed, as predicted earlier this Kosarev A.V.
with colleagues phonon flux responsible for the transfer of energy (including
the nerve impulse) in biomembranes [64 – 65]. In these works adequately
substantiated advantage of the phonon mechanism of transmission of nerve
impulses, compared with traditional ideas.
In my opinion, is more correct to use terms
such as "phonon package" or "phonon word" .
The main thing is the transmission of nerve impulses – not just the energy
transfer as such but information. Naturally, that information should be some way
to code, that being "recognized", adopted by the receiver, causing it
to one or another answer. That's why we put those terms to reflect the essence
of the signal – information. I.e. information signal – it's not just energy
transfer, but transfer the encoded and may say otherwise, that is the transfer
of the quantized energy.
Schematically, I tried to depict this process in
Fig. 9. It should be noted that I was trying to portray a quasiparticle, or
rather their totality, realizing the hopelessness of the process. It should be
noted that the process of the terminology necessary for describing the MMB
has just begun, as the study began and the phonon nature of nerve impulses, it
is possible in the future will be proposed and justified for other terms, but
for now enjoy these.
Inactive protein
Protein complex
"Phonon package"
with an external signal
"Phonon word"
Fig. 9.
An attempt to schematically show the formation of
"Phonon-pack" or "phonon-word". (Explanations
are given in the text below)
Under the influence of an external signal
membrane protein carries out some process, such as transmembrane ion channel
opens, changing thus the conformation (structure). Such sharp changes in the
protein structure causing a disturbance in the surrounding areas near the
protein in the hydrocarbon chains of lipid molecules. They get a "hit"
that leads to biophonon. For simplicity, and "clarity"
deliberately omitted other possible processes that have been described in the
above equations and discussed in previously published articles.
In such a system usually has to occur an
information signal, possibly consisting of more than one phonon, and possibly
having a 3-dimensional structure. This follows from purely steric (spatial)
parameters of membrane molecules. It is obvious that such a complex phonon
signal carries much more information than that used in conventional electronic
computers, one-bit signal. That is why the terms were introduced –
"phonon package" or "phonon word". The signal
can be received by the addressee, in whole or in part. Also, the sender can send
multiple identical or different signals to different destinations. Then, the
sender under the influence of surface tension of the biomembrane ΔÐ
returns to its original state. Membrane protein
system is
again ready
for use.
Schematically, this process
is shown in Fig. 10.
The most trivial variant of the process described
is shown in Fig. 10. In response to an external signal, structural rearrangement
occurs in the molecule, which increases the molecule diameter; that is, the
molecule passes into the excited state (Fig. 10 A). Increasing of the molecule
diameter is accompanied with the work against the membrane lateral compression
forces. So, the protein activation energy transition into the energy of
surrounding membrane lateral compression occurs. Such energy dissipation brings
about heating of the membrane (in the classical approximation this process is
described by the equation PV = RT), hence, of the cell. Then,
after accomplishment of the elementary act of signal transformation, the protein,
under the influence of the membrane lateral pressure, returns to the initial
state (Fig. 10 B). Several other similar schemes could be suggested, however,
this one should suffice to understand the principle.
If we now consider Fig. 10B, it becomes
evident that black triangles are quasi “loudspeakers” through which the
protein emits the surplus energy of excitation in the course of relaxation into
the initial state, and this energy is emitted as a phonon. In its turn, the
phonon can be scattered on the defects of the quasi-solid bilayer fragment, its
energy being channeled to heating the cell as a whole. Besides, the phonon can
propagate along the quasi-solid bilayer fragment to the next protein membrane
system (either an individual molecule or an ensemble of various molecules) and
serve for the latter an external signal. In Fig. 1 A this would be depicted as
another black triangle (but this time, a phonon receiver) at the non-excited
protein molecule. The arrow “signal” would be shifted to the membrane so as
to point to this triangle. It could coincide or appear close to the arrow of
“relaxation” in Fig. 10
B. Further details are evident from this scheme.
It is now possible only express the most general
considerations about the specific mechanism of transmission of intracellular
information using biophonons. Have it is obtained that there is an
intracellular language, words of which are "phonon packets."
This is all information that is known about the cellular language for this
moment. But it may be important for the problem of understanding the mechanism
of "biological computer". In our case, may be said that more
specifically, the information relates to the biomechanical machines [66],
because we consider the biomembranes, as the classical and quantum mechanical
systems. In other words, it is about understanding the possible mechanisms of
transmission of information that a section of a cell subjected to external or
internal influence, the coordination of the various membrane and/or
intracellular protein systems, etc.
Now it becomes clear the role of the disordered
hydrocarbon phase of the lipid molecules in the center of the lipid bilayer of
the biomembrane. Because of the disordered structure, biophonons may not
be distributed in the central region of the biomembrane. Thus, there is a
separation of information flows in biological membranes. Consequently, the need
to have nonlipid molecules for signals transmission from the outside of the
biomembrane and vice versa. Thus, the transmembrane protein of the system must
perform one more function – to provide information exchange between the inner
contents of the cell and its surroundings. Now you can with a certain pain safe
to say that the boundary between the living cell and its environment takes place
in the central region of the lipid bilayer of biological membranes.
Quantitative studies of the transformation and
transmission of information signals, the mechanism of their effect on the life
of cells, questions related to the work of cell signaling systems are beginning
to engage more and more of the various scientists and research groups around the
world [67 – 73]. The authors note a number of advantages biocomputers what can
be seen live cell systems, compared with those that we usually use: a colossal
memory capacity, micro sizes, versatility, etc. Much more important is the study
of cellular systems with multiple states. Moreover, the transitions between
these states are provided by control inputs of different nature.
Here's an example: suppose that the cell has
sixteen different states and, furthermore, it interacts with many of the same
neighboring cells. Of course, a cell with sixteen states by itself enough to fit
that, but associated with the billions of congeners, it can form a memory
unthinkable capacity. Figuratively speaking, a teaspoon of the solution with the
bacteria could have the memory of all the current computers. But the question
arises how to program such biosystems?
The problem, at least, complex, and on the way to
her researchers have to solve another one - to learn how to manage the processes
of interactions between proteins and genes. How to enter and display information
from the bio-computers? I do not seem promising to manage such biocomputer by
chemical reactions [73, 74]. So it is very slow and difficult way – it is
unlikely that he realized in nature as the sole and universal way.
However, the biophonons mechanism
of interaction membrane systems first theoretically predicted and experimentally
confirmed makes a simple and understandable process of information exchange
between different membrane systems, interference on one another and synchronizes
their work.
Fig 10.
The Simplest scheme of the membrane proteins operation. (The
Explanation in text.)
As part of MMB may be explained, and
many aspects of the functioning of the nervous system process transmission of
nerve impulses becomes a simple, clear and consistent [5 – 8, 10, 29 –
31, 75, 76].
Briefly the essence of the process lies in
the fact that the receptor neuron, receiving an external signal (stimulus) is
activated, emitting "phonon package", utilizing one or more C-atoms of
the lipid hydrocarbon chains. (Fig. 9, 10)
This "phonon packet", then spreads to the ordered
regions of the biomembrane. Later, perhaps, that regardless of
me the nature sounds a nerve impulse arrived by other authors
[77]. However, they suggest that the nervous impulse is distributed in the form
of solitons – structurally stable solitary waves propagating in a nonlinear
medium [78]. But it is difficult to speak of a stable wave, consisting of one or
two phonons. A sound wave is still a close-up characteristic of the environment
and the processes occurring in it. Hopefully, it's just unfortunate use of the
term. If we put aside because the soliton, a term related to the macrosystem,
and appeal to his sense, then here it coincides with the previously introduced
the terms "phonon word" and/or "phonon package." Just have
to reiterate that while in MMB no generally accepted terminology, and MMB
itself is not yet there.
First of all, let us estimate the order of
the time required for biophonon pass one lipid molecules in biomembranes.
Phonons propagate speed is equal to sound speed in this medium, which for liquid
oils is ~ 1000 m/sec. Linear size of the lipid molecules in biomembranes
with their characteristic ΔÐ
is equal to ~ 10 Ǻ. Then the time required to pass a single phonon
lipid molecules ~ 10-10 sec. This time several orders of
magnitude smaller than the characteristic time describing the motion of lipid
molecules colliding as a whole in biomembranes, which was mentioned earlier in
this paper.
Thus, for the time necessary to overcome
the phonon distance of several cm lipids in biological membranes can be regarded
as virtually immobile. This means that biophonon
distributed in nearly regular, crystalline structure
characteristic of the polar regions of the biomembrane. However, although the
membrane for periods of time ~ 10-10 – 10-9 sec
and can be viewed as biocrystal, in reality it has the properties of the gas
condensed, but the gas. It should present some defects in the form of the
density fluctuations, shifts the lipid hydrocarbon chains relative to each other,
etc. In such time intervals that are characteristic of the phonon signal for
distribution in biological membranes, the latter can be regarded as an imperfect
crystal containing defects in their structure. The presence of defects leads to
distortions in the structure of the transmitted information signal, weakening it
– a reduction of its intensity and the emergence of other noise. Therefore,
when sending a signal to the "long distance" course at the cell scale,
it is necessary to periodically increase and adjust, i.e., clean up and correct
the possible distortions in the signal.
The need to transmit data signal to the
"long range", sometimes several tens of centimeters, there is in the
axons of nerve cells (in neurons). In the works related to the study of alcohol
and other organic solvents and other compounds soluble in the hydrophobic region
of biological membranes, it is shown that they violate the transfer in the
membranes of bio-phonon data signals through the creation of a bilayer lipid
matrix defects in its structure, and possibly at the expense of changes its
compressibility. The internal volume of the neuron is used for life support of
the cell. There the intracellular transport of those components, synthesis,
metabolism of proteins, neurotransmitters [79] and other molecules necessary for
cell function as a living system. And most importantly, ensures the functioning
of the cell "information" of molecular membrane system, maintaining
its efficiency and constant readiness to transmit signals, as transmission of
information and transport of substances needed
for the cell space divided among them. (Simplified description,
but does not alter the fundamental picture.)
In any textbook on information transmission
may be read to signal that the need to periodically adjust its increase and its
possible distortion. Therefore needed intermediate repeater station, which may,
if necessary, to produce more, switching and/or transformation (encoding), they
received signals. These stations must be in constant readiness and be provided
with independent power source. In our case we are dealing with a "molecular
amplifier", which must be periodically placed along the length of the
neutron (the axon). The structure of the axon satisfies all these requirements.
We see a regular structure; along which there are equal
nodes of Ranvier (intercepts). Then
we get the information signal, and the accompanying potential jump that occurs
in these areas – well known to all potential actions that accompany the
propagation of nerve signals along the axon, and the resulting work of molecular
"amplifiers". Naturally for the molecular phonon amplifier used source
of energy that is always "in hands"
– a source of electrical energy, is always present in biological cells –
the transmembrane potential difference. It does not matter which particular
point or points of eclectic capacitor, what is the lipid bilayer of the
biomembrane, capacity, because on the plates (plates) of the capacitor, it is
installed everywhere the same. Membrane potential exists all over cell surface
and may be used at any point and at any time. It does not really matter where
the molecular "electric generator" and the speed of his work (his
power). The slow speed of the "generator" may be compensated for their
number. The only thing that it is desirable to do – to isolate the membrane
surface to compensate its spontaneous discharge. The myelin sheath of axons that
way it looks and has all the necessary properties: it is a good electrical
insulator, and also prevents the penetration of biological membranes of
surface-active compounds that may cause the formation of structural defects in
the lipid bilayer matrix. In other words, in this case, we have all the
necessary and sufficient for the functioning of molecular quantum-phonon
membrane information chain.
It is obvious that to destroy the molecular
system
possible in several ways: to destabilize the signal
transduction pathways and/or disrupt the molecular "amplifiers".
Enough to break down one of the "molecular amplifier" on the line, so
all the line is not working. It needs to recover, or seek ways around to
transfer the information. Of course, that the simultaneous violation of
signaling pathways and molecular amplifiers, which may be added signal sources
and receivers of these signals is finite, will reach its goal – the
destruction of the whole information system of cells – faster and have more
varied. In other words, it is obvious that the larger units and pieces of cell
information system we break, then the more destabilizing the whole system work.
It seems that some drugs – opiates, cocaine and
others – are acting specific to the protein part of the "molecular
amplifier", blocking the activity of individual sections of membrane
proteins. Drugs temporarily block the work of membrane amplifiers, and
neurotoxins bind irreversibly, completely blocking the transmission of nerve
impulses.
There are non-specific compounds which may "dissolved"
in the lipid bilayer regions of biological membranes. One of the best known
compounds of this kind is alcohol (C2H5OH) with the action
that is facing a wide range of people, not just experts who study it especially
from a professional point of view.
It is reasonable to assume that the better
solubility of the molecules in lipid membranes, then, respectively, all other
things being equal, they would form a greater number of defects and stronger
influence on the parameters of the membrane, the stronger the activity of these
compounds. I .e.
it is obtained
by previously
known Meyer-Overton
rule [80].
Must again be noted that alcohol, organic
solvents, anesthetics and other similar molecules of different nature, including
the gases used for anesthesia, within the framework MMB have fundamentally
similar molecular mechanism of action. The mechanism of action depends on their
solubility in membranes, rather than on the chemical structure of the compounds.
That is why the description is carried out and an alternative mechanism for this
rather broad class of compounds. Of course, in between there may be differences,
but these differences are not fundamental, which also coincides with the
findings obtained in independent studies [29 – 31].
All above stated can be formulated briefly as
follows: foreign molecule "dilute" the inner hydrocarbon region of
lipid bilayer of biological membranes, disrupting the functioning of membrane
proteins, preventing their activation and/or return (relaxation) to its original
state once they complete the elementary act of its operation. This occurs
because change the integral parameters of biomembranes – its lateral surface
pressure (surface free energy of the interface membrane - washing her solution)
and the lateral compressibility. In this same process can be violated exchange
of signals between different protein molecules (systems) within the biological
membrane. What is especially important for the signals in nerve cells. More
details on this process considered previously [29 – 31].
The action of these compounds (alcohol and other
organic "solvents") leads to the fact that the cell perceives their
presence, as the action of high temperature, which also gives rise to defects in
the quasi-crystalline matrix of the biomembrane. This is the thermal defects
that arise due to increased thermal motion of molecules. But this rise in
temperature only "apparent" real organism lives in the same conditions
that had lived before, but these conditions are perceived as abnormal (hotter
than they really are).
Nervous system exists that would quickly
send information signals from the periphery to the brain for processing and
issuing commands to the executive organs and tissues, for example, to perform
any movement by the muscles.
Naturally, this is a quick transfer of the
received signal to the addressee - it is the whole system exists, which requires
expenditure of energy from protein system –
the amplifier. Often, the speed and accuracy of the
transmission and analysis of information received depends on the very life of
the organism. As a result, inactivation of this system – dominoes fell (Fig.
6). Then the system needs to be translated to its original state - you need to
again raise and set dominoes. There is already another
form of transformation needed energy (chemical and/or electricity) for which the
most likely to have effects other protein systems (or subsystems). Electrical
membrane potential should be used (this is logical) in the transfer of the
phonon signal "molecular amplifier" because it exists on the entire
surface of the biomembrane, but not ruled out the use of other energy sources.
However, in the case of chemical energy must be targeted to deliver the required
connection to the place of their use. And this is a relatively slow process,
which is the limiting stage of diffusion of compounds within the cell along a
gradient of electrochemical potential. At this stage, returning to its original
state – may well be relatively slow, perhaps running through a few steps or
intermediate states. The slow reaction is easily offset by their number. If the
return of molecular amplifiers in the initial state depends on the transmembrane
potential, it is natural that, affecting the latter, we will disrupt the entire
nervous system. And it's fairly well-known process electro anesthesia
[78, 79]. To do so, impose the electrodes on the head with general anesthesia or
to a particular nerve trunk innervating the area for which you want to process
pain. The best effect is achieved by applying to the electrodes of rectangular
pulses with a frequency of 100 – 200 Hz. It is possible that this frequency is
associated with the time needed to bring the molecular amplifiers in working
condition after the operation, i.e. perform a one-time act to strengthen and/or
correction of the transmitted signal. Obviously, this is sufficient gross
interference in the membrane of molecular amplifiers can lead to their
inadvertent operation, resulting in spontaneous twitching of muscles under the
influence of an external electric potential, which is not always acceptable.
The mechanism described suggests the existence
and other physical methods of anesthesia, which will no longer have the
disadvantages of the previously existing methods and/or upgrade the last of
these same goals.
Thus, it appears that the recorded neurons
upon excitation the action potential is not by nerve impulses, but the result of
recovery after the operation of the amplifier of the membrane to its original
state ready for use, i.e. to strengthen and/or correction of the transmitted
information signal photon. In other words, that uses pre-stored cell energy of
the electric field that is available anywhere at
the cell membrane.
It is shown that the phonon mechanism of
transmission of nerve impulses well explain the existence of threshold values,
when in response to stimulation of the nerve impulse does not occur
[65].
In conclusion, of this
part of article gives picture of (Fig. 11), where the author
also tried to portray realistically the non-existent quasi particle – a phonon
[80].
Fig. 11 .
Another attempt to portray a phonon.
How could the author cope with
this task – to judge you, but this picture is similar to that and I was trying
to portray.
closing
Stated in this article
MMB allow us to consider biological membranes as a mechanical system obeying to
the same time and in one the same place the laws of classical and quantum
mechanics. The boundary between quantum and classical effects is the velocity
occurring in living systems processes. Quanta that transmit the
mechanical interaction in biomembranes are (bio) phonons that not only
transfer energy from one system to another protein but may be while information
signals of still large enough information capacity – "phonon package"
or "phonon word" acting in the living systems.
In the beginning was the Word,
and the Word was with God…
In Him was life…. (John
1.1, 1.4).
Thus, the membrane possible to speak,
are a mixture of two quasi gases condensed two-dimensional quasi
crystal gas variable curvature consisting from molecules and gas,
non-material quasi-particles – bio phonons.
Both are characterized by gas-specific parameters for each cell, the first gas (membrane
lipid matrix) – to its surface pressure and compressibility, and the second
– Σ ħωi – their frequency and structure.
These parameters of both gases allow existing and function in biological
membranes are not arbitrary molecules, but only those that correspond
to these parameters. This fact is significant for pharmacology
and genetic engineering. Not any molecule working in
the membrane its native cells can
penetrate and operate in the membrane of another cell.
Part of the other conclusions stemming from
the MMB, were considered in the text of this article and references cited.
This conclusion concerning the mechanism of action of alcohol, local anesthetics,
sound, etc. It
is described the mechanism extreme gas disinfection of
various equipment, volumes,
air-conditioning systems, etc. in situations where the object is not known and/or
no time for it identification.
Really in this article
it is impossible to describe all the conclusions arising from
the MMB. Purpose of the article was a popular exposition of the basic
concepts and rationale of the terms used an explanation of their physical nature.
In conclusion I would like to briefly
mention some of the medico-biological aspects that are specific to MMB.
Conducted the initial measurement of the phonon spectra of various living cells,
showed their suitability for identification of the latter [7, 81], and moreover
many-component systems. Naturally, the normal and abnormal cells are different
from each other. It is not difficult to work on one type of cells without
affecting other cells in the system. The
simplest effect on cells is the initiation of their death, that just as suitable
for the diagnosis and
control of tumors.
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Publishing date: September 28, 2011
Source: SciTecLibrary.ru
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