ISSN 1990-7478, Biochemistry (Moscow) Supplement Series A: Membrane and Cell Biology, 2009, Vol. 3, No. 2, pp. 101–115. © Pleiades Publishing, Ltd., 2009.
Original Russian Text © F.I. Ataullakhanov, N.O. Korunova, I.S. Spiridonov, I.O. Pivovarov, N.V. Kalyagina, M.V. Martinov, 2009, published in Biologicheskie Membrany, 2009,
Vol. 26, No. 3, pp. 163–179.
101
1 The ionic asymmetry of cells, i.e., the fact that the
concentrations of Na
+
, K
+
, Ca
2+
ions inside the cells are
strongly different from their concentrations in extracel-
lular liquids, is known for a long time [1]. This phe-
nomenon has been and is explained in different ways
[2]. It seems quite natural to assume that such asymme-
try provides the sensitivity of cells to damage [3–5].
Another supposition, indisputable with respect to excit-
able cells, is the statement that the ionic asymmetry is
needed for electric potential generation and for ensur-
ing the electric activity of cells [6–9]. One more version
is that the presence of ion gradient permits active trans-
port of various useful substances into a cell [10–12].
Each of these assumptions is reasonable in relation to
the same types of cells and may be disputed in relation
to other types. However, ionic asymmetry exists in all
cells without exception. Moreover, the concentrations
of univalent cations are practically the same in the cells
of all types [1].
Such surprising uniformity compels us to think that
the ionic asymmetry is an element of one of the most
basic cell systems. In recent years is has become clear
that it is really so, because any cell consists of a great
number of molecules, which are absent in the environ-
ment, and these molecules generate an excess osmotic
pressure in a cell, so that cells must resist osmosis and
control their volume. This is the same basic problem for
1
The article is translated by the authors.
any cell as, for example, energy supply. For the first
time, the idea of association of ionic asymmetry with
osmoregulation was apparently suggested in the work
of Tosteson published in 1959 [13]; it was described in
more details by Tosteson and Hoffman in 1960 [14].
Nearly half a century has passed since that time, and we
finally begin to understand that the necessity of main-
taining constant cell volume appears simultaneously
with the very first cells in the course of evolution. For
this purpose, nature has invented ion pumps, special
channels, and a number of other notable systems, which
provide the cell with fantastic stability and autonomy.
We are still far from complete understanding of the
whole picture; however, what we know anyway opens
up a harmonious and beautiful multilevel system of cell
volume control. This regulation has been studied best in
the red blood cells (RBCs) of mammals; hence, volume
control in a human RBC will be in the focus of attention
in this paper.
Mathematical models played a key role in the study
of cell volume control [15–21]. The models provided
answer to quite a number of questions, which were
either very difficult or impossible to answer experimen-
tally, predicted a series of unexpected connections in
cell metabolism, and helped us to understand why a cell
needs some senseless (on the face of it) biochemical
reactions.
In recent years, mathematical modeling has become
popular. At the same time, the overwhelming majority
How Erythrocyte Volume Is Regulated,
or What Mathematical Models Can and Cannot Do for Biology
1
F. I. Ataullakhanov
a–c
, N. O. Korunova
b
, I. S. Spiridonov
b
, I. O. Pivovarov
c
,
N. V. Kalyagina
d
, and M. V. Martinov
b
a
Center for Theoretical Problems of Physicochemical Pharmacology, Russian Academy of Sciences, Moscow, 119991 Russia;
e-mail: fazly@hc.comcor.ru
b
National Research Center for Hematology, Russian Academy of Medical Sciences, Moscow, 125167 Russia
c
Physics Department, Moscow Lomonosov State University, Moscow, 119991 Russia
d
Bauman State Technical University, Moscow, 105005 Russia;
Received January 31, 2009
Abstract—Modern concepts of the red blood cell (RBC) volume regulation are considered. It is shown that the
system of ion pumps and channels in the cell membrane ensures the physiological value of volume with a pre-
cision of about 10% even at 5- to 7-fold variations of passive membrane permeability for ions. Particular atten-
tion is paid to mathematical models for evaluation of the role of different molecular mechanisms in the RBC
volume control. It is shown that many questions, for example, ‘why the Na
+
,K
+
-ATPase pumps the ions in oppo-
site directions’ or ‘what is the physiological role of Ca
2+
-activated K
+
-channels’, cannot be answered without
adequate mathematical models of such complex control systems as cell volume control.
Key words: red blood cell, Na
+
,K
+
-pump, Ca
2+
-activated K
+
-channels, cell volume, mathematical models.
DOI: 10.1134/S1990747809020019