F = force; R =distance 
dependen ,ce of potential energy; d = distance between two 
particles. Vander Waals' forces are both (a) long-range at· 
tractive and (b) short-range repulsive. The relationship of 
force F and distanced is expressed as the resultant force c, 
i.e., a+b, represented by the curve c:k/tP, where k is a con­
stant that depends on the particles. At any distance shorter 
tlun d, the repulsive force increases mpidly, which prevents 
maner from collapsing. that when values of d are less than those for wruch 
U ltfJ is :zero, the U AB rapidly increases in positive 
value. This positive potential energy function counter· 
acts the possibility of collapse of matter. The principle is illustrated in Fig. V111: I, showing 
the well known distribution curve of resultant poten­
tial energy (c), composed of attractive (a) and repulsive 
(b) forces. Such forces vary experimentally and theo­
retically for different kinds of matter and their degree 
of purit}• (4). For instance, different kinds of carbon 
(such as coal, graphite, and diamond) present a wide 
range of internal cohesion. Fixed angulation or rota· 
tion of particles also influence the resulting forces. In a system of many particles it is assumed that most 
but not all of the concentration forces are pairwise 
additive (3). Among the three types of so-called long-range 
forces, e/ectroscacic forces (I) are either repulsive or 
attractive. They also are pairwise additive according to 
the principle of superposit ion (simple addition) of the 
fields. Electrostatic forces dominate large separations. In the case of dipole-induced dipoles (II), however, the 
forces are not pairwise additive. They arise by the 
interactions of the permanent electrical moments of 
76 Concentration-dispersion forces particles. The long-range dispersion forces (Ill) are ex­
clusively attractive and pairwise additive. These forces 
are also called c®centration forces (9, 10, II). Thus, long-range for·ccs acting upon several parti­
cles can be written as a sum of the individual electrical 
forces and dispersion (concentration) forces, when 
consideration is given to the spatial orientations and 
rotational moments of the participating particles. Derivation of the contributions of the long-range 
forces to U AB have been given by Hirschfelder, et al. (6), as follows: 
I) The long-range electrostatic forces may be written 
as a series 
U electrostatic= VN•+ UJlQ+Uw+ ... 
where each partial term is a potential energy function 
of separation, orientati.on, as well as electrical mo­
ments of the particles (see also Fig. X: 2). Up,u repre­
sents the potential eneTgy function of the electrical 
charges J1 + and Jl-of a IJJI dipole. It may be described 
by the fu.nction (p2/d3) f, where f is a function of 
angulation and d the distance between the two charges. UpQ is a similar function between the neighbouring 
and attracting opposite charges of the I'Jl dipole and 
another dipole of charges Q+ and Q-. U00 is the 
similar function for the QQ dipole. Likewise, all other 
combinations of dipoles in the given collection of parti· 
cles are added into this electrostatic series. The values 
of U!lQ and UQQ are much smaller than UpJl. They 
fall off, according to Margenau (II), proportionally to 
d-• and d-5, respectively, whereas U,up is propor­
tional to d-3. II) The long-range inducrio11 forces produce balanced 
net forces between panicles without additive effect on 
surrounding matter (3). Their importance for structur­
al development within a given field should therefore 
depend to a large extent on the dielectric properties of 
the material in which induction takes place. III) The third type of long-range forces, the concen­
tration forces (dispersion, London-van der Waals, co­
hesive adsorption forces), are believed to arise as a 
specific dipole effect on a particle by electron move­
ment in an adjacent particle. Thus London (9, 10) has 
shown that the overall energy function Ud;sp may be 
written as a series of exclusively additive terms propor­
tinal to -kld6 for each pair of particles. The concen­
tration forces can be determined but require know­
ledge of the particular wave function for ground and 
excited states of the electrons. Formulas for their cal­
culation have been developed (8, 10, 13, 15). The counterba lance of the negative long-range at­
tractive forces is the repulsive short-range van der 
Waals' forces. These forces rapidly increase their 
dominance within distances of approximately 3 A. 
Tills effect depends on the increasing repulsion caused 
by t'verlap of the electronic charge clouds of particles as they move closer to each other (II, 12). This repul­
sive potential ener«Y is described by a term propor­
tional to t""' or d-• where p and n are constants (n 
varies between 9 to 24) (3). 1\ potenual tuncuon lor overlap repulsion between 
water molecules was described by Kamb (7) in the 
form 
u,.pu, =A" ( ~ )'. where o•276x 10-1 em ("collision diameter'' of the 
molecules ). Two pairs of coefficients and exponents 
were derived: A*=2.1 kcal molufe-1, n= 10 and A •= 
4.0 kcal mol-1, ns9 for ice of IWO different energy 
contents. Interatomic forces and short-range molecular forces 
are known to be integrated into the combined or hy­
brid movements of orbital electrons around atoms of a 
molecule (3, 4). Two electrons move with opposite 
spins in each orbit around the molecule, producing 
internal bonds of the molecule within certain sections. The variations in electron density of certain sections of 
the orbits allow interactions between adjacent mole­
cules. When external forces have vanished, this muru­
al process produces an equilibrium configuration of 
the participating nuclei. The short-range repulsive 
forces must then be balanced by an electrostatic attrac­
tive charge. This attraction is limited to the binding 
regions of the molecule, whereas all other regions tend 
to separate the molecules. A specific molecular orien­
tation takes place at the binding sites. The balance of 
the forces involved in the binding follow the Hell­
mann-Feynman rule (3, 4), which states that the 
forces acting on the nucleus of a separated molecule 
are the sum of the electrostatic forces arising from the 
other nuclei and the force derived from electronic 
charge density. In the discussion on physicochemical potential, the 
chemical potential contains the activity component a1• 
The innuence of this factor is not simply a maner of 
concentration of j. The "total activity" of solute j is 
related to both iu concentration and other energy 
components, which are included as an activity coeffi­
cient, y1: 
The activity coefficient is usually synonymou s with the 
electric characteristics of solute j and differs for 
charged particles in water solution by their electrical 
charge from the activity being caused only by the 
concentration. This variation was recognized by De­
bye and Hiickel (I) who also found that, as the concen­
tration increases, the electrical fields of the ions inter­
fere increasingl y with each other. Consequently, local 
accumulations of a species of ions of varying concen­
tration will interact different internally and with sur-rounding reactants. The activity facton for ions in 
solution also vary by their different net electric charge 
in relation to the activity of surrounding media. In 
arldition, thermal factors, pressures , gravity and exter­
nal electric fields will, of course, also inlluence ionic 
dispersion . In a local region around any ion, the elec­
tric forces influence the movement of other ions. As 
concentration increases, the distance between ions de­
creases, which facilitates ion-ion interactions . The presented view of ionic potential has its corre­
spondence in the molecular potential of certain bio­
logically active nonionic compounds. Associated prob­
lems are particularly considered in the presentation of 
Ll-te ionar~rgonar concept of energy exchange over 
biologically closed electric circuits (Chapter XIII), an 
important prerequisite for the utilization of their bio­
logically stored electrical energy. This c.olldttued survey of some clraraeteris1ics of molecu­
lar and particulatt forces of attraction and repulsion may 
an jim imprmion umr far distant from problems of 
srrucrural rrwdif ~eation in biological materia!. II is not so. These forces constitute importam portia/ functions in a 
complicattd systtm of many intmuring forces, all of which 
must be considertd in order to understand rek.ttd problems. Some of these functions will next be shown in stud­
i<S of movement of water in organized tissue (Chapter 
IX) and then of the behaviour of particles, i.e., nonor­
ganized corpuscular clements (Chapter X). Gross morphologic development of tissue is compli­
cated particularly by the superimposition of both \'llri­
:.able and more stable intenactins components. Among 
these components, convection, diffusion, and tissue 
matrices constitute important fa.ctors. To this group 
belongs also a hitherto overlooked, important compo­
nent: biologically closed electric circuits. References 
t. Dcby<, I. P., and Huct..l, E.: Zur Th«<i< ckr El<kuolyte I. 
Phyo. Zeitschr. H: 185, 19Z3. 2. Eds.tll, j. T., and Wyman, J.: Cit. in BiophysK:al chemistr)', 
Vnl. I. New Vnrk. Academic Prel>.<;. l9SA. 3. Eisenberg, 0., and Kauzmutn , W.: The: llfUCtuR and propc:r· 
ria of ... , ... Oxford, Oarenclon l'r<u, t969. 4. Feynman, R. P., Leighton, R. B .. and Sands, M.: The Feyn­
man lectures on physics. New York, Addison-Wesley Publ. Co., t%4. S. Hamaku, H. C.: The London-Van dcr Ww auroction be­
twcc,n t:pherial panicles . Phyliea 4: 1051, 1931. 6. Hinchfelder,J. 0., Curtiss, C. F., and Bird, R. 8.: Mokculor 
theory o( ga!tS 1nd liquids. N~ York, \l1ilt}', 1954. 1. Kamb, 8.: In: Ri<h, A., and Oavicbon , N. (cds. ): Structwti 
chemistry and mol«Ub.r biology. San Fra.nasco. Freeman, 
1968. 8. Klrkwood, J. G.: The dielectric polarlzatlon of polar liquids. J. 
Chcm. Ph)'$. 7:91 t, 1939. 9. London, F.: Zur Tbeorie und SySlnnatik der }\o\ekulari:riftc . Z. Phyo. 63: 24S. 1930. Concentration-dispersion forces n 10. London, F.: The gtneraJ theory of molecular forces. Trans. Fa .. day Soc. 33:8, 1937. 11. Margenau, H.: Van der Waals forces. Rev. Modern PhysiC$ 
II: I, 1939. 12. Margenau, H., and Myers, V. W.: The forces between water 
molecules and the second \'irial coefficien t for water. Phys. Rev. 66:707, 1944. 13. Muller, N.: Concerning structural models for water and chemi­
cal-shift data. J. Chem. Pbys. 43:2555, 1965. 78 Concentration-dispers ion forces 14. Rowlinson, J. S.: The second vidal coefficients of polar gases. Trans. Faraday Soc. 45:974, 1949. 15. Slater, j. C.: lnuoduction to chemical physics. New York. McGraw-Hill, 1939. 16. Stockmayer, W. H.: Second virial ooefficien ts of polar pses. Chem. Phys. 9:398, 1941. IX. Water 
E lectroosmotic transport over closed electric circuits 
In Chapters III-V it is indicated that tumours in tbe 
lung may produce local accumulation of fluid (a "B" 
zone) between the pleura and the lesion, due to block­
ing of draining pathways between the pleura and the 
hilum. The fluid accumulation in the lung is often 
seen separated from the surface of the tumour by 
another zone, wruch shows radiograpruc signs of de­
crease in fl.uid content (an "A" zone). The existence of 
the two zones may be explained as a result of local 
lymphoedema produced by stasis and an electroosmo­
tic process. Electroosmotic transport of water in tissue 
will be described in this chapter as a function of 
biologically closed electric circuits (BCEC). A. Movement of water into 
necrotic tissue 
Early stages of autolysis of a tissue include increased 
weight of the tissue and accumulation in it of calcium 
(21, 32). Conway and McCormack (18) showed that 
the freezing point depression of autolyzing tissue in­
creases rapidly, indicating an increased osmotic pres­
sure. Furthermore, direct measurements of ionic con­
centrations in injured tissue indicate that the increased weight and swelling are caused mainly by a net uptake 
of water, sodium and chloride (45, 59). When hypoxia induces injury of tissue, proteolytic 
enzymes are released (32). Molecular degradations en­
sue, thereby increasing the osmotic properties of the 
injured tissue (63). Water then enters the tissue. Lo­
caUy increased fluid pressure should also contribute to 
a further progression of the necrosis. Water content of mammalian tissues may also in­
crease during hypoxia, without permanent damage to 
cells (2, 81). It has been suggested (63) that the intra­
cellular fluids are hypertonic and that respiration gov­
erns water balance of tissue because energy is expend­
ed for active transport of water out of the cells. Increased content of water in necrotic neoplasms 
can also be expected for other reasons. Consider the 
case of fixed negative charges in the matrix of a tu­
mour barrier (defined as the viable tissue surrounding 
a necrotic centre) and the presence of a closed electric 
circuit containing a gradiem of electric potential over 
the barrier. Electroosmotic flux of water should then 
take place toward the cathodic tissue. Such a biologi­
cally closed electric circuit (BCEC) and its activation 
will be described in Chapters XII and XIII. For the 
moment, we will assume only that BCEC systems exist 
and investigate the consequences of such a mechanism 
on transport of water in tissue. In the case a tumour is 
Water 79 electropositive in relation to the surrounding tissue, an 
outward flow of water, away from the tumour, should 
take place. This effect should then be detectable radio­
graphically as a hydropenic "A" zone and the dis­
placed water, if locally accumulated around a tumour, 
as a hydropic "B" zone. Conventional radiographic 
techniques, however, are not capable of identifying a 
local increase in water content inside a tumour with a 
cathodic centre. B. Intercellular space and 
movement of water through 
ussue 
An important effect of forces of attraction and repul­
sion in tissue is connected with the existence of inter­
cellular spaces. These forces balance each other, leav­
ing a distance d open (Fig. Vlll: 1). The rapidly in­
creasing component of repulsion force therefore pre­
vents adjacent cells from achieving "absolute con· 
tact". It is easy to understand that the space d should be 
of considerable importance for the functions of cells 
and tissues. Physically, a space must be present be­
tween individual cells. Biologicall y, a space sufficient­
ly large to allow water and solutes to pass between the 
cells is an obvious prerequisite for cellular functions . Current belief is that the adhesive bonds generally 
maintain intercellular distances of no greater than 5 A 
and that the repulsive forces maintain intercellular 
distances of no less than 3 A (7, 15, 20). The size of a 
water molecule has been found to be 0.9572±0.0003A 
(7, 8). The vibrational state does affect the dimensions 
of the water molecule, but only slightly (22, 40). Pas­
sage of water molecules between cells is therefore 
physically possible. The spaced is indeed a space of life. A further possibility is that passage of water mole­
cules between cells may open further the intercellular 
space for new molecules, the attraction forces between 
adjacent cells having lessened. Whatever the mechan­
isms may be to modify the space d, it is obviously a 
large enough space to permit the entrance of water. C. Fixed surface charges 
on cells 
Bernstein (9, 10) proposed in 1902 the "membrane 
theory" on the basis of the Nernst-Planck concept of 
the liquid junction potential (see Chapter Xlll). He 
80 Water thereby identified fundamental physical characteristics 
of molecules and cells. Diffusion potentials across 
membranes and electrical phenomena of living cells 
and tissue became to a certain extent explainable. Bethe and Toporoff (I I) next pointed out, in osmo­
tic studies, that the direction of deviation of the con­
centration potential from the liquid junction potential 
correlates with the electrokinetic charge of a mem­
brane. Teorell (82, 88) formulated in the 1950's his 
"ftxed" charge theory, that ionic membranes have 
two Donn:~~n equilibria and one net diffusion potential. Meyer and Bemfeld, (57) in particular, and later many 
others (17, 24, 37, 38, 43, 44, 55, 56, 66-69, 71-74, 
77, 79, 80, 93) have further extended the basic concept 
of fixed charges of membrane s and their many impor­
tant functions in biology. For example, fixed surface 
charges are considered to exist on the surface of both 
plant and animal cells (I, 36, 90-92). Large molecules 
of organic substances containing a polar part are often 
arranged periodically in so-called pseudocrystallinic 
aggregates (15). Such aggregates are present in cellu­
lose, wood, silk and cotton wool, as well as in many 
animal proteins (3, 29). Some of these aggregates are 
in the form of surface-coating lipoproteins with a mi­
celle structure. The polar parts of these molecules at 
the cell surfaces are thought to act as ftxed surface 
charges (27). The water-absorbing properties of many 
substances are also considered to depend on the ftxed 
charges of the absorbing material. Cotton wool, which 
has a large surface per weight, may absorb as much 
water as 25 times its weight (3). The surfaces of cells are known to carry fixed elec­
tropositive and electronegative charges (92). A surplus 
of negative charges almost always characterizes the 
surfaces of cells (1, 92). Two adjoining cells electro­
statically attract each other at certain sites where the 
flXed charges have opposite polarity (1, 4, 36, 38, 50, 
75, 91). These charges are important for the mechani­
cal strength of tissue. Fixed surplus charges are also 
important constituents in the structural prerequisite s 
for transmembranous and regional transport of water 
in interstitialtissues(61, 75, 94). D. Liquid water: structure 
and energy 
Before we can discuss electroosmosis adjacent to 
lung lesions, some physical properties of water must 
be reviewed (26, 42, 44, 62, 77). The structural fea­
tures of liquid water depend largely on its ability to 
form bonds by combinat ions of the molecular orbits of 
electrons of its hydrogen and oxygen atoms. Liquid water also appears in polymers that possess a varying 
degree of vibrational characteristics (7, 8). In ice, representing the lowest energy level of water, 
hydrogen -bonded clusters of water molecules form te­
trahedrons (26). As energy content in liquid water 
increases, the numbers of tetracoordinated bonds de­
crease. Thus triple, double and unbonded water mole­
cules appear progressively as energy increases to the 
level of vapour, which represents the highest level of 
energy content (60, 75). The difference in energy be­
rween the levels for the tetrrabonded and unbonded 
water molecules is estimated to be 2. 7 kcal/mole. Brownian and vibrational moveme nts of water mole­
cules constitute an important part of their total free 
energy content. Because a water molecuJe possesses a permanent 
dipole moment, it will orien.t itself within an electric 
field, i.e., liquid water is "structured" in a way similar 
tO what takes place in the formation of ice (60). An 
augmentation of the dipole moment will also develop 
when the molecule is exposed to an electric field. E. Electroosmosis: transport 
mechanisms Types I-IV 
Tbe mode of water transport in electroosmosis is 
known to a certain extent (44, 70, 77). Transport of water by use of fixed charges on cellu­
lar surfaces can be demonstrated in the following ex­
periment (Fig. IX: 1). Cotton wool is packed in the 
lower part of a U-shaped glass tube, forming "capilla­
ries". A matrix now exists of spaces or channels among 
the cotton fibres. Water is poured into the tube. Two 
clean platinum electrodes are immersed in the water. When an electric potential is applied between the elec­
trodes, water moves from the positive to the negative 
side of the system until an equilibrium is obtained 
berween the electroosmotic and hydrostatic pressures. This flow of water, against a hydrostatic pressure, may 
be regarded as a primitive form of "active transport" 
(defined as transport against an energy gradient), for 
which the presence of suitably sized capillaries lined 
with fixed charges is just as essential as the applied 
elt-ctric field. Such transport of warer (electroosmotic 
water transport of what may be called Type I) can be 
assumed to take place without concomitant electrolysis 
of waler molecules . In a simplified version, Fig. IX: 2 
illustrates this electroosmotic transport of Type I. 
I. Type I electroosmosi .s 
When an applied electric field is strong enough to 
break up clustered water mol.ecules, a field orientatio n 
or "structuring" of the molecules wiU take place. 7-814$86 NordtJUtrOm + 
Fig. IX: I. Electric transport of water. Conon wool is 
packed in the lower part of aU-shaped glass tube, which is 
partly filled with water. Two clean platinum electrodes are 
immersed in the water. When an electric potential difference 
is applied between the electrodes , water is moved in lhe 
.. capiUariesu of the wool from the c:lcctropositivc to the eJcc· 
tronegativc side of the system. An equilibrium is obtained 
between opposing ele<:troosmotic and hydrostati c forces. Simultaneously, the molecules and dielectric compo­
nents of the matrix will increase their dipole moments. The dimensions of the matrix of "capillaries" are also 
important. Lined with fixed charges, the capillaries 
must be smaU enough to prevent hydrostatic return. Under these conditions, water will move in the capil­
laries in a predictable way. Negative surface charges of the capillaries will push 
some of the oriented molecules (Fig. IX: 2, site X) and 
puU some of them (site Y). Vibrational and rotational 
movements of the water molecules can still take place 
in irregular capillary channels (77). Certain molecules 
will be transported along the matrix secondary to the 
motion of the molecules in sites X and Y. The pres­
ence or absence of net electrostatic forces on a mole­
cule of water in the matrix largely depends on the 
Water 81 Type of 
electro-+ 
osmosisr------------..---------~~--------~------------~ 
X y 
I t+ Dielectric -J H, ••••••••••••••• H-FfO ANOMAL. OSM .• H-0 I+ a = = = :.1 • 
n ... • NET ELECTROPHORETIC 
H20 TRANSPORT 
Base Acid 
m I CHARGED POLYMERS •• 
+ 
N I <£H.0 DIPOLE INDUCTION <0R21 
Fig. IX: 2. Electroosmotic water transport. four mecha­
nisms can be identified. Type I transport requires fixed elec­
tric charges on the surfaces of the dielectric matrix (e.g., cot­
ton wool). These charges are negative in cotton wool and tis­
sue, leading to a net movement of water molecules from an­
ode to cathode. The energy for this l1'1lnspon is suggested 10 
deri\'C from the applied c:lcctric field transforming the: energy 
of the water molecules. The transformations include break­
ing of clustered wate.r molecules, and reduction of their vi­
brational and rotational molecular movements. A net move­
ment of water is produced by electrostatic interaction be­
tween the die-lectric matrix and aligned water molecules. Type I I transport is electrophoretically linked and requires a 
closed electric circuit but no fixed surface charges on the d.i-
location of the fixed negative charges in relation 10 the 
actual charge distribution of !he water molecules. Dur­
ing transport in the matrix, !he dipoles of the molecule 
and the fiXed charges are exposed 10 synchronously 
fluctuating forces from their electrical fields, depend­
ing on their spatial locations in relation 10 each other. A different dipole moment of !he field-oriented water 
molecules is also to be expected depending on their 
location in !he positive or negative side of the superim­
posed electric field. The direction of water' flow in electroosmosis is 
determined by the polarity of the surplus of fixed 
82 Water elecuic "capillary'' walls. When water is ionized, the differ­
ent speeds of migration of ions and their recombination cre­
ate a net transport of water. Type Ill transport is cationic. It 
is based on the fact that cations adsorb water molecules , but 
anions do not. The cations therefore selectively carry water 
molecules in a closed circui1. This type also includes l1'1lns­
port of ionic pol)•mc.rs of watc.r. Type IV electroosmosis (in­
duction-adsorption ) takes place in m31rices close 10 an elec­
trode and does not require ftxed surface charges on the "cap­
illary'' walls. Water molecules undergoing dipole induction 
by the electric field are transported (adsorbed) to an adjacent 
charged electrode- surface, irrespective of its polarity. The 
"capillaries" of the matrix may be thought of as mechanical 
rectifiers. charges of !he capillaries (C), within a defined c.lectric 
field. Sollner (77) studied electroosmosis through a typical 
unoxidized collodium matrix membrane of high poros­
ity. Thereafter he oxidized !he same membrane with 
heparin, which is electronegative , and prepared an-other membrane with protamine, which is electroposi­
tive. An electroosmotic flow was then obtained for a 
10-3 M KCl-water solution (current intensity, 0.1 
mA/cm2) as follows: 
Unoxidized collodium 
membrane 
Oxidized collodium 
membrane 
Protamine collodium 
membrane +4640 mm3/IOO cm2/hour 
The rate of transport of water depends on several 
factors: magnitude of the electric field over the pores, 
the membrane porosity (number and size of pores), 
and the magnitude , density and geometry of fixed 
charges. As voltage increases over the electrodes, 
movement of water will increase. A similar effect can 
be obtained by placing the electrodes closer to the 
matrix, due to the increase of intensity of the electric 
field. The size of the "capillary" channels also may be 
critical. If the channels are too large, hydrostatic pres­
sure will cause water to return. This effect may reduce 
the difference between the fluid levels or may even 
lead to no difference at all. 2. Type II electroosmo sis 
The mechanisms of "electroosmotic water transport" 
are complex. It is easy, for example, to recognize that 
water can also be transported as a result of electrolysis 
followed by migration of protons and hydroxyl ions 
and their recombination. In such instances semiper­
meable membranes or capillary channels need not be 
charged (Fig. IX: 2). The membranes or channels then 
act simply as a mechanical hindrance to convection. In Type II electroosmosis, diffusion and electro­
phoretic migration of participating ions in an applied 
electric field determine where the recombined water 
molecules will be found. The transport mechanism of 
the migration derives from the mobility of ions. Proton 
movement, for instance, is defmed as 36.2 · w-• cm2 
v-• s -• at strong dilutions and 25°C; the corre­
sponding value for hydroxyl ions is 20.7 · 10-4 cm2 
v-• s-• (41). Their distance of recombin ation into 
36.2 water can then be expected to be 36.2 + 20.7 or about 
¥1 of the interelectrode distance from the electroposi­
tive electrode. This value is, however, also modified 
by different adsorption propertie s of ions in a ntatrix 
as well as by gravitation. (We will here refer to Fig. XIII: 7 which, besides serving a different purpose, also 
illustrates Type II electroosmosis. ) In this experiment a 
10 volt DC potential was applied between two plati­
num electrodes on litmus paper soaked in water. By 
electrolysis of water, protons are produced at the an­
ode and hydroxyl ions at the cathode. The ions mi-grate in the electric field and recombine to form wa­
ter. Due to differences of mobility of the ions, a net 
transport of water will occur toward the cathode (see 
Fig. IX: 2). Gravity considerab ly influences the posi­
tion of the recombination zone, an effect which can 
easily be observed by slight tilting of the electrolytic 
specimen . Water transport of this kind (electroosmo­
sis, Type II) increases in importance as voltage in­
creases and is, in fact, a special case of electrophoresis. In the conventional view of electrophoresis, transport 
of electrons in the electrical cables requires a corre­
sponding electrical transport i n the water. This trans­
port has been explained as depending on proton jumps 
on water molecules (26), after their ionization at the 
electrode surfaces. However, electrolysis of water can 
be induced immediately on applying a potential differ­
ence over water, despite the fact that water is a good 
electrically insulating medium. Some dissociatio n of 
water is evidently present for an initial ionic flow of 
current, but other mechanisms may also be involved. Dielectric induction and structuring of water mole­
cules adjacent to the electrode surfaces can, for in­
stance, be regarded as the initial prerequisite for redox 
reactions at the electrode surfaces. Water molecules 
are thereby oriented in an optimal way to consume 
electrons from the cathode and to donate electrons to 
the anode. The reaction products then diffuse and 
migrate in the electric field. This mechanism is com­
patible with the relatively slow transport of OH-and 
H+ ions in the direction from the electrode surfaces to 
the zone of recombination, which should be a barrier 
of resistance for the passage of protons and hydroxyl 
ions. 3. Type III electroosmosis 
Another of the mechanisms of electroosmotic water 
transport depends on differences in behaviour of ca­
tions and anions (Fig. IX: 2). It is known that cations 
may become hydrate-d , but not anions (41). When ions 
migrate in the electropositive part of an electric field, 
cations should then carry water molecules adsorbed on 
their surfaces. A correspond ing adsorption and trans­
port by anions should not occur in the opposite direc­
tion. The resulting net electrophoretic water transport 
might then he looked on as a primitive form of "medi­
ated"' transport. A release of transported water mole­
cules from the carrier can even he predicted when the 
aggre,gate enters a region of high pH. 4. Type IV electroosmosis 
Finally, an electroosmot ic partial function may he dis­
tinguished (Fig. IX: 2). Within a matrix, which docs 
not need to be lined with fixed charges, strong dipoles 
Water 83 Fig. IX: 3. Appantusfor circularelectroosmotictransportof 
watet. The electroosmotic chamber consists of two perforat· 
ed plastic (perspex) plates. Cotton wool is compressed be· 
tween the plates. On the other side of each plate is a plati· 
num electrode and a venical tube. The tops of the vertical 
tubes are connected by a slightly tilted tube. The apparatus 
is filled with water up to the level of the lower end of the tilt­
ed connector. Positive voltage is applied to the right elec­
uode and negative t.O the left one. Electroosmoric Oow of wa­
ter will then take place from the right to the left chamber. Increased hydrostatic pressure of the elevated left column of 
water will be limited by the flow of water from the left to the 
right vertical tube over the slightly tilted tube. The circular 
movement of water will then be seen as a dripping of water 
from the "mouth" of the tilted tube. This flow will continue 
for a while. Then it will slow and stop spontaneously , de­
sphe mainttna.nce of t,he voltage over the electrodes. Fig. /X:4. Pressures in electroosmotic transport of water: 
The tihed connecting tube in the apparatus shown in Fig. IX: 3 has been replaced by a pressure transducer in the left 
vertical tube. Upon application of a 20 volt DC potential he­
tween the electrodes (left negative, right positive) a rapid rise 
in pressure (phase a) was followed by a slow rise (phase b). Two minutes later, when the electric potential generator was 
turned off, the pressure rapidly diminished (phase<). Phase 
a is thought mainly to be caused by Type I electroosmosis, 
and phase b by continued cJecuophorctic water transport 
(mainly Types II and D1 electroosmosi s). The remaining 
pressure (d minus c) is produced by gas formed in the left 
chamber. The electrophoretic water transport during phase 
b is caused by pressure c minus a. 
c 
d a 
84 Water may be induced in the molecu.les close to a charged 
electrode. These inductions will cause a net attraction 
force (adsorption) of water to any charged electrode, 
regardless of its polarity. The matrix in this Type IV 
electroosmosis serves then as a rectifier for the trans­
ports. F. Two distinguishable 
pressure variables in 
electroosmotic transport 
of water 
One critical question is whether appreciable electroos ­
motic water transport can occur at voltages low enough 
not to create electrolysis of water. Different opinions 
have been presented in the literature on this problem 
(76). Helmholtz (39) found electrolysis of water to stan at 
about 1.64 V. Bartoli (6) found the value to be 1.23 V. 
Part of the movement of water during electrolysis in 
electroosmosis is then connected with the formation of 
H~O+ and its polymers (corresponding to Type Ill 
electroosmosis ). In search of further insight into the 
mechanisms of electroosmosis, the behaviour of water 
in terms of transport pressures was assessed experi­
mentally. l. Experime ntal methods and results 
An electrophoretic apparatus was constructed as 
shown in Fig. IX: 3. It consists of a chamber and two 
platinum electrodes, each attached to electrical cables. Between the electrodes, various materials can be 
pl~ced under compressio n between two perspex plates 
with multiple holes. Vertical tubes are connected with 
the right and left pans of the apparatus. A slightly 
tilted tube connects the tops of the vertical tubes. In 
the arrangement shown in Fig. IX: 3, the electro­
phoretic matrix consists of compressed cotton wool. When, for instance, 10 volts arc applied between the 
platinum electrodes, the right one positive and the left 
negative, water will flow through the cotton from right 
tO left. This flow results in a return of water from the 
left to the right side through the slightly tilted tube, 
appearing as dripping of water from the mouth of the 
tilted rube. The degree of compression of the cotton wool, the 
distance between the electrodes , and the potential dif­
ference each were found to influence the transport of 
water. These influences were seen as differences in 
numbers of drops of water per urtit time. The rate of 
water transport was not found ro be constant. It 1mmHp 
volts 20 18 16 14 12 10 9 8 7 6 5 4 3 2 
Fir.IX: S. Pressures in electroos.motic trans-oort of water 
when fourteen DC pulses of one second each were applied. The rapid pressure phase returned to the initial level of pres-
reaches a maximum after some time, depending on 
voltage, then decreases and eventually stops complete­
ly. This change in water transport is associated with 
formation of gas at the electrodes, which also elevates 
the free fluid levels in both vertical tubes. The tem­
perature of the water will simultaneously rise, the 
more rapidly as voltage is elevated. After transport of 
water has stopped, renewed transport can be obtained 
by elevation of the electric potential between the elec­
trodes. The water will then start to flow with increas­
ing speed, again reaching a maximum and then retard­
ing until it again stops. These prclhuinary observations were fun her studied 
by removing the tilted communicating tube and con­
necting a pressure transducer (Statham P23 DC) to the 
left or right vertical tube, which was closed by an air­
tight stopper of low compress ibility. The apparatus 
was ftlled with water, replacing all air pockets. The 
pressures were recorded by means of a Grass 7 B Poly­
graph. When a constant potential of 20 V was applied 
berween the electrodes for 2 minutes, rwo components 
appeared in the changing pressures obtained from the 
electronegative side of the system (Fig. IX: 4). Initial­
ly, pressure increased rapidly (phase a) and then it 
rose slowly and steadily (phase b). When the voltage 
was switched to zero, the pressure diminished immedi­
ately (phase c), which is larger than the initial elevation 
of pressure (a). The new base level of the pressure was 
eh:valt:d, at the value d r:t•inu:s , .. In the next experimental variant, the values of (a) 
and (c) were the same during a consecutive series of 14 
applications of square voltage pulses of one second 
duration each (Fig. IX: 5). The voltage pulses varied 
from 20 down to 2 V (Grass S88 stimulator). The 
magnitudes of the initial a pressures were directly 
proportional to the applied voltages (Fig. IX: 6). sure after each of these short pulses. indicating that remain­
ing effects of ionization and gas formation must have been 
minimal. Intermittent applications of20 V for 2 minutes each, 
separated only by turning tbe applied voltages off and 
then immediately on, gave results which are shown in 
Fig. IX: 7. The slow phase now steadily built up the 
pressure in the system. Ionization causing the forma­
tion of gas, visible on the electrodes as bubbles, is the 
probable main source of this increase. The insert in 
Fig. IX: 7 shows the increasing total pressure (tl) of the 
system. At the same time, the rapid phase (a) also 
shows increasing values with each repeated application 
of 20 V. This effect is possibly a consequence of 
increasing ionization of water, which should relatively 
enhance Type Ill electroosmosis. Fig. IX: 6. Electroosmotic transport of water: hydrostatic 
pressures increase during a series of one second pulses of in· 
creasing voltage. The pressures were proportional to the po­
t·cntial differences between the electrodes (voltages ranged 
between 0.4 and 20 volts). The relationship suggests linear 
correlation. mm 
H20 
1.5 
lO 
0.5 
.. 
0o~----~5~----~ t~0----~~~5--­
volts 
Water 85 mmH20 
8 
Fig. IX: 7. Electroosmotic uansporr of water: pressures in· 
crease during intermittent applications of 20 volts for periods 
of2 minutes each. The slow pressure phases caused the pres· 
sure in the system to increase continually , an effect due 
mainly to gas, forming as a consequence of ionization. Such 
2. Discussion of pressure changes 
and electric transport of water 
It is now apparent that rwo separate phases of pressure 
changes are integrated in electroosmot ic water trans· 
port. The slow, cqntinuous pressure phase is obviously con· 
nected with ionization wd the formation of gas in the 
system. This phase is not easily recognized at low 
voltages with pulses of up to 20 V of I sec duration. Gas formation and consequently also ionization are not 
particularly prominent with this mode of application 
of voltage. The rapid pressure phase is easily observable at low 
and high voltages and immediatel y on the application 
of a voltage difference between the electrodes. Voltage 
pulses between 0.4 to 20 V and of I sec duration each 
have shown linear correlation with the pressures, im· 
plying that they may be extrapolated to zero. These 
findings have not been described previously. The gradual increase in magnitude of the rapid 
pressure phase on repeated applications of voltage 
shows that this phase also is partly coupled to the 
mechanism of the slow continual pressure phase and 
most likely to Type III electroosmosis. In the author's opinion, the rapid pressure phase in 
these experiments corresponds mainly to the transport 
process which is some·times called anomalous elec­
troosmosis (77), while the slow and rapid pressure 
phases are both included in what is usually called 
electroosmosis. It seems questionable to make a dis· 
tinction between anomalous osmosis and electroosmo­
sis, because each includes all four types of transport 
mechanisms. The internal relative importance of the 
Type I-III mechanisms for net transport of water 
changes as ionization increases. According to the pres· 
86 Water events can also explain the progressive increases of the rapid 
pressure phases (a). The insert shows the relation at two 
minute intervals (n) between the total pressures (If) in the 
system and the rapid pressure phases (a). sure measurements, the rapid pressure phase is an 
instantaneo us electric field effect which is most clearly 
d~monstrated upon application of short pulses of volt· 
age. The "low" voltages, e.g., 10-20 V between the 
electrodes, are "high" compared with what can usually 
be expected in biologic situations. Still, extrapolation 
of the rapid pressure phase with short voltage pulse 
suggests that electroosmosis also can occur over very 
low voltages between the electrodes. This observation 
is of interest, because it suggests that small gradients 
of potential should also be able to include transport of 
water. The amount of water transported may even be 
large when the forces are allowed to work over a long 
time. A more critical problem is that proof has not yet 
been put forward, demonstrating in tissues any chan· 
nelizing mechanism which could function as a closed 
electric circuit in vivo. Such a circuit is a prerequisite 
for electroosmosis. Chapters XII and XIII are devoted 
to this problem. So-called anomalous osmosis, known since its first 
description by Dutrochet in 1835 (25), has been much 
studied (5, 34, 35, 46-51, 70, 78). Graham suggested 
that anomalous osmosis is a special kind of osmosis 
(33). Bartell and Loeb (5, 52-54) have suggested that 
spontaneous electroosmosis occurs whenever electro­
lytes, but not non-electrolytes, diffuse across a charged 
membrane. A double layer of high field strength on the surface 
of cells, including vascular endothelium, is normally 
about 106 to 10" volts per centimetre (64). These 
electric fields are caused by fixed cellular charges 
which correspond in vivo to the fixed surface charges 
in vitro in the cotton wool experiments. The magni· 
tude and direction of anomalous osmosis directly de-pend on the fixed charges of membranes (77). De­
pending on the polarity of these charges, as well as the 
concentration differences across the membrane, 
anomalous osmosis may be positive or negative. In the author's opinion, all transport of water via a 
closed electric circuit is electroosmosis . This transport 
comprises four integrated but different mechanisms 
(Fig. IX: 2). These may be characterized as follows: 
Electroosmosis of Type I seems to be almost synony­
mous with previous descriptions of "anomalous (elec­
tro) osmosis". ft is characterized by its dependence on 
a surplus of fixed negative or positive charges in "cap­
illaries". Type I electroosmosis may therefore also be 
described as fixed charge electroosmosis. Its field-in­
duced rapid change of pressure correlates directly with 
the magnitude of the voltage applied over the matrix. Electroosmosis Type II does not require fixed charges 
in the matrix. It involves electrolysis of water, diffu­
sion and migration of H+ and OH-ions and their 
recombination into water as a special case of electro­
phoretic transport. Type II electroosmosis is charac­
terized by the slow elevation of pressure, which is not 
directly proportional to the magnitude of the voltage 
applied over the matrix. Type II electroosmosis may 
also be described as elecrroosmosis by ionic recombina­
tion. Type Ill electroosmosis is closely related to Type n 
electroosmosis. Its mechanism is based on ionic hydra­
tion (77) but applies only to cations. Cations therefore 
carry water and water polymers from the electroposi ­
tive to the electronegative part of the electric field. This function may therefore be looked on as a primi­
tive form of mediated transport. Type III electroosmo­
sis may also be described as cationic electroosmosis. Lastly, a Type IV electroosmosis may be recognized . ft takes place in a matrix close to charged electrodes , 
which by induction produce a net attractive force on 
nearby water molecules. In Type IV, no fixed charges 
are necessary in the matrix. The water molecules will 
always move to the adjacent charged electrode regard­
less of its polarity. Type IV electroosmosis may also be 
described as field-induced osmosis and should be regard­
ed as a special case of Type I electroosmosis. In Type 
IV electroosmos is the charged electrodes include the 
functions of both the applied field and the flxed 
charges. G. Transport energy in 
Type I electroosmosis 
The driving force in anomalou s osmosis (fixed charge 
electroosmo sis) through cell membranes, according to 
Sollner (77), may possibly be the streaming potential which occurs as an electrolyte diffuses. According to 
this view, the streaming potential difference as a 
source for the driving force should require a closed 
circuit, which may exist but is not yet demonstrated. Other authors have suggested that the whole clectroos­
motic process can be explained as an electrophoretic 
IJ'ansport of water (47-49, 58). This latter explanation 
is evidently oversimplied, because now two different 
phenomena of elec1ric transport of water can clearly be 
distinguis hed by their pressure characteristics. The energy of electroosmosis of the fiXed charge 
type may possibly derive from the thermal energy of 
water. This consideration merits discussion. According to Wagman eta!. (89), and Cottrell (19), 
the following energies are associated with the forma­
tion of a molecule of water: 
kcal mol-' 
(I) Energy of formation from atoms 
at 0 K -219.34 
(2) Zero-point vibrational energy 13.25 
(3) Electronic binding energy 
= (1) minus (2) -232.59 
(4) Enthalpy of formation at 2s•c -22.54 
(5) Bond energy of 0-H bond at 
0 K = Y2x(l) 109.7 
(6) Dissociation energy of H-0 lOI.S 
(7) Dissociation energy of 
H-OH = (I) minus (6) -117.8 
The free molecular energy of nonhydrolyzed water 
molecules is stored as vibrational and rotational move· 
ments. This energy could be available for Type I 
electroosmosis as follows: an electric field can be an­
ticipated to restrict molecular movements. This re­
striction will lead either to (a) loss of energy to the 
surroundings, or (b) modification of kinetic energy 
from the previous vibrational and rotational move­
ments into directed transport of the molecules, or (c) a 
combination of both possibilities. The breaking of clusters of bonded water molecules 
(16, 30, 31) and their "structuring" by an electric field 
means that the amount of "available" free energy of 
the water is modified in some way. This energy, which 
is part of the total energy content of water, cannot 
spontaneously return to its initial equilibrium state 
unless the electric field is removed. Some reactions 
toward an increase of entropy are still possible, but 
toward a different stage of equilibrium, which in the 
presence of fixed surface charges in capillaries may 
lead to transcapillary · electroosmot ic transport of Type 
I. The restriction of brownian movements of water 
molecules by an applied electric field might, theoreti­
cally, be reflected in a temperature change of the 
water. The effects of different water temperature on 
electroosmotic transports as well as different electrode 
Water 87 Fig.IX:8. Electroosmotic transport of water: temperature 
effects. Te.mperarure and pressure were recorded in the ap· 
paratus for electric water transport, immersed in a thermo· 
stat (C)·controlled water bath. A sensitive (0.01•C= 1 mm 
deflection) electro thermometer (E) was placed in the cotton 
wool matrix (.t\1). Matrix temperature was measured against 
a reference elecuothermome ter in ice water in a thermos 
flask (7). One second pulses of 10 to 40volts were applied 
(G) over the marrix at equilibrated matrix temperatures 
against the thermostat water in n~nges from + 1•c to +SO'C. The djsplacements of water were del'eroJned over the pres .. 
sure transducer (P). potentials on water temperature were therefore studied 
(Fig. IX: 8). The temperature of water inside the matrix of "cap­
illaries" (cotton wool) was measured with a sensitive 
(±O.oJ•C) electrothermometer (Philips Chrome! Alu­
mel Thermocoax ). The electroosmotic apparatus was 
immersed in a thermostatically controlled water bath. A sensitive pressure transducer (Statham P23 DC) was 
connected to the branch of the electronegative side 
while a mercury thermometer for rough controls was 
inserted into the open opposite branch. The electro­
thermometer was positioned with its tip in the cotton 
wool matrix and its reference electrode in a thermos 
flask containing ice water. Single electric potential 
pulses of one second duration were produced by a 
square pulse generator. Pressure changes in the elec­
tronegative side of the system were amplified and 
recorded (Grass Polygraph Model 7B). When stable temperature levels were applied to the 
external bath and measured inside and outside the 
apparatus, variations from + I°C to +500C did not 
influence the electroosmotic pressure elevations as po­
tential pulses of 10 to 40 volts of I sec duration each 
were applied over the electrodes . 88 Water At the same time, no elevation or depression of 
temperature was observed within the cotton wool ma­
trix. The range of water temperature in these experi­
ments, + I°C to +so•c, includes most of the actual 
range of biologic temperatures. The absence of detect­
able influence of water of different temperatures on 
electroosmotic pressure as well as the absence of de­
tectable temperature changes within the cotton wool 
matrix during electroosmosis suggest two pos.sible ex· 
planations. First, the sensitivity of the pressure and 
temperature recording devices may have been too low, 
which is not very likely. Second and more likely, the 
transformation of the available free energy of water 
may correlate directly with the energy level of the 
applied electric field. This possibility is consistent 
with the previously described experimental results 
(Fig. IX: 6), showing a linear relation between applied 
electrode voltages and fiXed charged (Type I) elec­
troosmotic pressures. This explanation is in itself not 
particularly remarkable: isothermal reactions are com­
monly encountered in biology. H. Experimental 
electroosmosis in dog and 
human lung tissue 
Normal, fresh, peripheral lung tissue from 2 dogs and 
2 humans after pneumonectomy were each carefully 
minced. The tissue materials were then washed repeat­
edly in water. Each specimen was placed under slight + 
compression between the perforated perspex plates in 
the electroosmotic apparatus (Fig. IX: 9). The appara­
tus was then filled with water. Voltage was applied 
between the electrodes. Pressure was recorded in the 
two water chambers. These electroosmotic experiments revealed that hu­
man and dog lung tissue behaved in the same way as 
cotton wool. From these experiments it is apparent (Fig. IX: 9) 
that short pulses through a closed electric circuit can 
induce similar rapid pressure gradients over fresh hu­
man and dog lung tissue and cotton wool. Electroos­
motic water transport may possibly, therefore, also 
take place in the lung, e.g., as Type I electroosmosis 
( = fixed charge electroosmosis = "anomalous osmo­
sis"). Such a possibility , however, warrants some fur­
ther considerations. I. Electroosmotic flow of 
water: local displacement of 
water in the formation of 
"A" and "B" zones around 
a tumour 
Many studies have estimated the possible importan ce 
of electroosmosis in biology (14, 20, 23, 25, 26, 28, 34, 
35, 46, 65, 70, 77, 85). A small electroosmotic move­
ment of water toward the negative pole in the large 
cells of Nitella was found by Blinks and Airth (13, 14) 20V 
+ 
F~'g. IX: 9. Electrophoretic transport of water: matrix of 
freshly minced tissue from the lung of a dog. Pressure eleva· 
tion in the electronegative side (left upper) and correspond­
ing lowering in the electropositive s1de (left lower) of the ap­
paratus for electric water transport. Twenty volts for one 
second were applied across the electrodes. Reversed pres· 
sures were obtained when the polarity of the elecuodes was 
reve.r. ;ed (right upper and lower pulses). Dog and human 
lung tissue each appear to allow electric transport of water in 
the same way as does compressed CQnon wool. In this experi· 
ment traces of blood and elec!tolytes were present. Never­
Uu:le:o;:o;, the rapid pressun: pha~es wt:n: vf appruxirmud y llu: 
same magnimde as those with a matrix of deionized water 
and cotton wool (cf. Fig. IX: 5). but only at potential differences of J-2 V, which they 
considered to be 20 times greater than common bio­
logical levels. An exceedingly small electroosmotic flow has been 
demonstrated in Nitella by Penson and Dainty (28), 
but they concluded the flow was too small to be of 
physiological significance. 20 mm3 H20/coulomb was 
transported by electroosmos is. The physiological cur­
rent across the membrane by ion transfer is about 
w-' A/em', which should allow a water flow of only 2xw-• H20 em-' sec-1• 
According to these calculations, electroosmotic flow 
should have minimal if any significant function in 
biology. Further information in the literature suggests 
otherwise. S:.wycr ~ et al. (64), -state that transverse 
streaming potentials or pressure equivalent electroos­
mosis (65) exist as a result of pumping of positively 
charged ions and water (compare Type Ill, cationic 
electroosmosis ) through the negatively charged "Swiss 
cheese" pores of blood capillary walls. A pressure drop 
of as much as 100 nun Hg is then maintained across at 
most a few microns of intima. The electric double 
Water 89 Pleul'a r;:::========= =====::J! H[ITUu~s 
Oedema 
Oedema Eleetroo smotic 
fotce 
Fig. IX: 10. A tumour in the lung obstructs lymphatic flow 
toward the hilum by blocking some of the interlobular 
spaces. Oedema then develops in the tissue between the tu­
mour and the pleura. Electroosmosis will move water away 
from the tumour, in the presence of a closed electric circuit 
between an electropositive tumour surface and a relatively 
electronegative, surrounding pulmonary matrix of narrow 
channels lined with fixed negative charges. An equilibrium is: 
obl.ained between the electroosmotic pressure and [he elevat­
ed hydrostatic turgor pressure. The outward displacemen t of 
\\rater will produce a radiopaque "B" zone, which may also 
be called a hydropic zone. The displaced water will be re­
placed 'by increased quantity of air in the alveoli, contribut­
ing to the radiolucent (hydroponi c) "A" zone. layer at the vascular interface and at the surfaces of the 
cells within the walls presents a surprisingly high field 
streng~h. It approaches 106 to 108 volts per transmem­
branous em (64). As indicated by Teorell (83-86) and 
others, certain cellular functions can be explained by 
assunting elecuoosmotic transmembranous water 
transport. Thus, pressure-volume changes in cells 
might be coupled to transmembranous concentration 
changes of flow and to a postulated closed circuit 
electric transport over the cell membrane, producing 
electroosmotic flow. In this way an electrochemica l­
mecha.nical system could be described which may have 
a biological correspondence (87). The polarity of the net surface charge of cellular 
surfaces has been reponed to be almoSt exclusively 
electronegative (4, 12, 36, 92). One curious exception 
are spirochaetes, which carry a positive net surface 
charge (1). Previous investigators of electroosmosis in biology 
have dealt with these problems in connection with 
transmembranous water transport. In tihe present investigation s the possibility of inter­
stitial water transport is of particular interest. The 
basic physical requirements for such a mechanism are 
narrow channels with fixed charges, a closed electric 
circuit and a driving electric potential gradient. The network of intercellular spaces forming such 
90 Water channels ("capillaries ") in lung tissue appear to poS­
sess the capacity to produce electroosmosis, according 
to the in vitro experiments here presented. Because 
most mammalian cells are provided with a surplus of 
fixed negative surface charges, it can be anticipated 
that the electroosmotic water transport is partly medi­
ated by these surface charges of the cells, analogous to 
the case with cotton wool. This possible Type I 
mechanism of electroosmosis does not exclude the 
presence of other mechanisms of electroosmosis in 
tissue. The assumption that electroosmotic water transport 
is likely to produce "A" and "B" zones around differ­
ent lung lesions still requires an electric potential gra­
dient in a closed electric circuit. Chapter VII suggests 
that a possible energy source for activation of such a 
circuit is local degrading of tissue due to injury, bleed­
ing, local necrosis, infection, etc. Chapter XII will 
present the structure and Chapter XIII the principle of 
activation of such a closed circuit. This circuit will be 
seen to permit an understanding not only of electroos­
mosis in tissue but also of several aspects of structural 
development and function. For the time being, we may preliminarily assume 
that the prerequisites exist for electroosmotic water 
transport. Fig. IX: 10 illustrates some consequences in 
the lung. A tumour blocks the interstitial channels 
which normally carry fluid centrally from peripheral 
lung and the pleural space. Stasis results and fluid 
collects between the tumour and the pleura. Let us 
also assume that an electric field is superimposed be­
tween the electroposi tive tumour and the surrounding 
parenchyma and that the interstitial spaces are lined 
with a surplus of fixed negative charges. Furthermore, 
let us assume that a closed electric circuit exists be­
tween the tumour and the surrounding tissue. Given these prerequisites , water must move through 
the tissue away from the surface of the tumour. The 
hydrostatic (turgor) pressure and the electroosmotic 
pressure must then equilibrate. A hydropeni c "A" 
zone and a hydropic "B" zone will form. The author has attempted to determine the actual 
water content in the "A" and "Bn zones in human 
lungs after pneumonectomy and in animals post mor­
tem. Theseattcmpts have been unsuccessful in demon­
strating changes in water content in lung peripheral 
to tumours, despite radiologic demonstration in vivo 
of the two zones. This negative result is readily ex­
plained by the fact that the mechanical changes on 
opening the chest will modify the pressure conditions 
which allow tissue water to be retained in the hydropic 
zones. As has radiographica lly been shown earlier, the 
"B" zone, representing local accumulation of fluid in 
tissue, disappears immediately after the introduction of 
air into the pleural space. (In the female breast "A" and "B" zones also may develop around carcinomas. In such cases it has been 
possible to demonstrate a difference in water content 
between the "A" and "B" zones. These studies are 
reported in Chapter XVI.) Later we will return to tbe problem of local elec­
troosmotic water transports in vivo, in connection with 
therapeutic polarization of tissues (Chapter XVII). At 
this point, however, it is now appropriate to consider 
the corpuscular effects of chemical concentration 
forces and superimposed electric fields. References 
J. Abramson , H. W.: Electrophoresis of cells and proteins. New 
York, Hafner Publ. Co., 1968. 2. Aebi, H.: Elcktrolyt-Akkumulierun g und Osmortgulat ion in 
Gcwqssc:hnitten. Helv. Physiol. Acta 11:96, 1953. 3. Asthbury, W. T.: Fundame-nta ls uf fibre suucturt. Oxford, 
Univ. Press, 1933. 4. Bangham , A. D., and Pelh.ica, B. A.: The adhesi\•eness of cells 
and the natu.re of the chem.icaJ groups at their surfaces. Proc. Roy. Phys. Soc. Edinburgh 29:43, 1960. 5. Bartell, f. E.: Membran e potentials and their relation to 
anomalous osmosis. In: Mathews, J. H. (ed. ): Colloid symJ,XI· 
sium monography . Dept. Chem., Univ. Wisconsin, Madison, 
1: 120, 1923. 6. Banoli, A.: Ober die Zersetzung des Was.sen durch cine schwa. chere elecuomotorische Krafr als dir des Danidl'scben Ele­
mentes. Beibl. Ann. Physik 2: 566, 1978. 7. Benedickt, W. S., Gailar~ N., and Plyler, E. K.: The vibration­
rotation spectrum of D,O. j. Chern. Phys. 21: 1301, 19S3. 8. Benedickt, V/. S., Gailar, N., and Plyler, E. K.: Rotation· 
vibration spectra of deutented water vapor. J. Chern. Phys. 24:1139, J9S6. 9. Bernstein, j.: EJcctrobiologie. Bnunschwe ig, Friedr. Vieweg u. 
Soho, 1912. 10. Bernstein, j.: Zur elektrochentischen Grundlage der bioelek­
trischen Potentiate. Bioche:m. Z. 50:393, 1913. 11. Bethe, A., and Toporoff, T.: Obc:r elektrolytische Vergange an 
Diaphragmen . Z. Physik. Chern. 89:597, 19JS. 12. Bikcrman , J. j.: Physical surfaces. New York, Academic Press, 
1970. 13. Blinks, L. R.: The relations of bioelectric phenomena to ionic 
permeability and to metabolism in large: plant cdls. CSHSOB 
8: 204, 1940. 14. Blinks, L. R., and Airth, R. l.: Electroosmosis in Nitella. j. 
Gen. Physiol. 41:383, 19S7. 15. Brandenberger, E., and Epprccht, W.: ROntgcnographische 
Chemic:. Basic:, Birkhiuser, 1960. 16. Buijs, K., and Cboppin, G. R.: Near-infrared swdies of the 
structu.re of water. I. Pure water. J. Chern. Phys. 39:2035, 
1963. 11. Caplan, S. R., and Mikulccky , 0. C.: In Marinsky, j. A. (ed. ): 
Ion exchange:. Vol. I. New York, Dekker, 1966, p. l. 
18. Conway, E. j., and McCormack , J. 1.: The tol81 inten:ellular 
concentra tion of mammalia n tissues compared with that of the: 
intercellular flujd. j. Physiol. 1: 120, 1953. 19. CottreU, T. L.: The strengths of chemic-..J bonds. London, 
Dutterworths , 1958. 20. Dainty, j., Croghan, P. C., and Fenson, D. S.: Electroosmosis, 
with some applications to plant physiology. Can. j. Bot. 41:953, 1963. 21. Dawkins, M. J. R., Judah, J. D., and Rec:s, K. R.: Factors 
influencing the survival of lh•cr cells during autolysis. I. Path. Bact. 77: 251, 1959. 22. Dennison, D. M.: The infra-red spectra of pOiyatomic mole· 
cutes. Pan II. Rev. Mod. Phys. 12: 17S, 1940. 23. Dick, D. A. T.: Ccl1 water. London, Bu!lerwonh s, 1%6. 24. Dr&. )', S., and Sollnc:r, K.: A theory of dynamk polyionic 
polenrials across membranes of ideal ionic selectivity. Biochem. Biopbys. Acta 21: 126, 1956. 25. Dutrocbe t, R. J. H.: De l'endosmosc des acidc:s. Ann. Chim. Phys. 60:22, 183S. 26. Eisenberg, D., and Kauzmann, W.: Tbc structure: and proper­
ties of water. Oxford, Clarendon Prns, 1969. 27. Epprccht, W.: Grundlagen der Kristallstrukturbestimmu ng. In: S<hin2, H. R. (ed. ): L<hrbuch der Rom8endiasnostik . 6 
Au.O., Band I. Stuttgan , G. Thieme, 1965, p. 78. 28. Fe.nson, D. S., and Dainty, j.: Electroosmosis in Nitella. Can. J. Bot. 41:68S, 1963. 29. Frank, H. S., and Evans, M. W.: Free \'Oiume and entropy in 
condensed systems. J. Chern. Phys. 13: 507, 1945. 30. Fr:a.nk, H. S., and Wen-Yang, W.: lll.lon·solvent interaction. Structun J aspects of ion-solvent interaction in aque<Jus solu­
tio.ns: a suggested pi~turt ofwattr s!fu~lurt . Disc. Faraday Soe. 24: 133, 1957. 31. Fr.ank, H. S.: Covale-ncy in the hydrogen bond and the proper­
tics of water and ice. Proc. Roy. Soc. London A247:481, 19S8. 32. Gallagher, C. H., Judah, j. D., and Rees, K. R.: Enzyme 
chAnges during liver amolysis. j. Path. Bact. 72:247, 1956. 33. Graham, T.: VII. The Bakcrian Lecture. On osmotic fortt. Phil. Trans. Roy. Soc. London 144: 177, 18S4. 34. Grim, E., and SoUnc-r, K.: The contributions of normal and 
anomalous osmosis to the osmotic effects arising across charged 
membranes with solutions of electrolytes. J. Gen. Phys:iol. 40:887, 1957. 3S. Grim, E., and SoUner, K.: True anomalous osmosis in multi­
so[ute model membrane systtms. J. Gen. Phys.iol. 44:381, 
1916(). 36. Haydon, D. A.: The-surface-charge of ~us and some other 
small panicles as indi~ted by tlect.ropboresis. Biochim. Biophys. Acta S0:4S7, 1961. 37. Helfferich, F., and Ocker, H. 0.: lonen.austau scher·mem·· 
brane.n in bi-ionischen Systemcn. Z. Physik. Chern. 10:213, 
19~7. 38. Hc:Jffc-rich , F.: Ion exchange . Ne-w York, McGraw-HHl , 1962. 39. Helmholtz, H.: Srudien Ober elecuische Grenzsdlichtcn. Ann. Phys. Chern. 7:337, 1879. 40. Herzberg, G.: Molecular spec-tra and molecular structure. 2nd 
ed. New York, Van Nostrand , 1950. 41. Higg, G.: Alhniin och oorganisk kemi. 4th ed. Uppsala, Alm· 
qvjst & WikseiJ, 1966. 42. Kavanue, J. L.: Water and solute-water interactions. San Fran­
ci$00, Holden-Day, 1964. 43. Kedem, 0., and Katchalsky, A.: Pecrmeability or composite 
mcmbnoes . Trans. Faraday Soc. S9: 1918, 1963. 44. Lakshminanyanaith , N.: Transport phenomena in artificial 
mcmbnnes . Chern. Rev. 6S:49l, 1965. 45. leaf, A.: Mainttnance of concentration gradients and regula· 
do;n of cell \'Oiume. Ann. N.Y. Acad. Sci. 72:396, 1959. 46. loeb, j.: The: influence of e:lectrolyte:s on the tlect.rification and 
the rate: of diffusion of water through collodion membrann . J. 
Gen. Physiol. 1:717, 1919. 47. Loeb, J.: Influence of the concentration of electrolytes on the 
electrification and rhe ra.te of diffusion of water through coJJo­
dion membranes. j. Gen. Physiol. 2: 173, 1920. 48. Loeb, I.: On the cause of the: influence of ions on the rate of 
diffusion of wat~r through collodion me-mbranes , I. J. Ge,n. Physiol. 2:387, 1920. 49. Loeb, I.: On the cause of the influence or ions on the rate of 
diffulion of water through collodion membranes , 11. J. Gen. 
Physiol. 2:563, 1920. 50.
631928626-Biologically-Closed-Electric-Ci-Bjorn-Nordenstrom-pdf.txt
Summarize the following text section accurately and concisely. Maintain specific details and numerical information, and use anti-repetition mechanisms:

        F = force; R =distance 
dependen,ce of potential energy; d = distance between two 
particles. Vander Waals' forces are both (a) long-range at· 
tractive and (b) short-range repulsive. The relationship of 
force F and distanced is expressed as the resultant force c, 
i. e. , a+b, represented by the curve c:k/tP, where k is a con­
stant that depends on the particles. At any distance shorter 
tlun d, the repulsive force increases mpidly, which prevents 
maner from collapsing. that when values of d are less than those for wruch 
U ltfJ is :zero, the U AB rapidly increases in positive 
value. This positive potential energy function counter· 
acts the possibility of collapse of matter. The principle is illustrated in Fig. V111: I, showing 
the well known distribution curve of resultant poten­
tial energy (c), composed of attractive (a) and repulsive 
(b) forces. Such forces vary experimentally and theo­
retically for different kinds of matter and their degree 
of purit}• (4). For instance, different kinds of carbon 
(such as coal, graphite, and diamond) present a wide 
range of internal cohesion. Fixed angulation or rota· 
tion of particles also influence the resulting forces. In a system of many particles it is assumed that most 
but not all of the concentration forces are pairwise 
additive (3). Among the three types of so-called long-range 
forces, e/ectroscacic forces (I) are either repulsive or 
attractive. They also are pairwise additive according to 
the principle of superposit ion (simple addition) of the 
fields. Electrostatic forces dominate large separations. In the case of dipole-induced dipoles (II), however, the 
forces are not pairwise additive. They arise by the 
interactions of the permanent electrical moments of 
76 Concentration-dispersion forces particles. The long-range dispersion forces (Ill) are ex­
clusively attractive and pairwise additive. These forces 
are also called c®centration forces (9, 10, II). Thus, long-range for·ccs acting upon several parti­
cles can be written as a sum of the individual electrical 
forces and dispersion (concentration) forces, when 
consideration is given to the spatial orientations and 
rotational moments of the participating particles. Derivation of the contributions of the long-range 
forces to U AB have been given by Hirschfelder, et al. (6), as follows: 
I) The long-range electrostatic forces may be written 
as a series 
U electrostatic= VN•+ UJlQ+Uw+. where each partial term is a potential energy function 
of separation, orientati. on, as well as electrical mo­
ments of the particles (see also Fig. X: 2). Up,u repre­
sents the potential eneTgy function of the electrical 
charges J1 + and Jl-of a IJJI dipole. It may be described 
by the fu. nction (p2/d3) f, where f is a function of 
angulation and d the distance between the two charges. UpQ is a similar function between the neighbouring 
and attracting opposite charges of the I'Jl dipole and 
another dipole of charges Q+ and Q-. U00 is the 
similar function for the QQ dipole. Likewise, all other 
combinations of dipoles in the given collection of parti· 
cles are added into this electrostatic series. The values 
of U. lQ and UQQ are much smaller than UpJl. They 
fall off, according to Margenau (II), proportionally to 
d-• and d-5, respectively, whereas U,up is propor­
tional to d-3. II) The long-range inducrio11 forces produce balanced 
net forces between panicles without additive effect on 
surrounding matter (3). Their importance for structur­
al development within a given field should therefore 
depend to a large extent on the dielectric properties of 
the material in which induction takes place. III) The third type of long-range forces, the concen­
tration forces (dispersion, London-van der Waals, co­
hesive adsorption forces), are believed to arise as a 
specific dipole effect on a particle by electron move­
ment in an adjacent particle. Thus London (9, 10) has 
shown that the overall energy function Ud;sp may be 
written as a series of exclusively additive terms propor­
tinal to -kld6 for each pair of particles. The concen­
tration forces can be determined but require know­
ledge of the particular wave function for ground and 
excited states of the electrons. Formulas for their cal­
culation have been developed (8, 10, 13, 15). The counterba lance of the negative long-range at­
tractive forces is the repulsive short-range van der 
Waals' forces. These forces rapidly increase their 
dominance within distances of approximately 3 A. Tills effect depends on the increasing repulsion caused 
by t'verlap of the electronic charge clouds of particles as they move closer to each other (II, 12). This repul­
sive potential ener«Y is described by a term propor­
tional to t""' or d-• where p and n are constants (n 
varies between 9 to 24) (3). 1\ potenual tuncuon lor overlap repulsion between 
water molecules was described by Kamb (7) in the 
form 
u,. pu, =A" ( ~ )'. where o•276x 10-1 em ("collision diameter'' of the 
molecules ). Two pairs of coefficients and exponents 
were derived: A*=2. 1 kcal molufe-1, n= 10 and A •= 
4. 0 kcal mol-1, ns9 for ice of IWO different energy 
contents. Interatomic forces and short-range molecular forces 
are known to be integrated into the combined or hy­
brid movements of orbital electrons around atoms of a 
molecule (3, 4). Two electrons move with opposite 
spins in each orbit around the molecule, producing 
internal bonds of the molecule within certain sections. The variations in electron density of certain sections of 
the orbits allow interactions between adjacent mole­
cules. When external forces have vanished, this muru­
al process produces an equilibrium configuration of 
the participating nuclei. The short-range repulsive 
forces must then be balanced by an electrostatic attrac­
tive charge. This attraction is limited to the binding 
regions of the molecule, whereas all other regions tend 
to separate the molecules. A specific molecular orien­
tation takes place at the binding sites. The balance of 
the forces involved in the binding follow the Hell­
mann-Feynman rule (3, 4), which states that the 
forces acting on the nucleus of a separated molecule 
are the sum of the electrostatic forces arising from the 
other nuclei and the force derived from electronic 
charge density. In the discussion on physicochemical potential, the 
chemical potential contains the activity component a1• 
The innuence of this factor is not simply a maner of 
concentration of j. The "total activity" of solute j is 
related to both iu concentration and other energy 
components, which are included as an activity coeffi­
cient, y1: 
The activity coefficient is usually synonymou s with the 
electric characteristics of solute j and differs for 
charged particles in water solution by their electrical 
charge from the activity being caused only by the 
concentration. This variation was recognized by De­
bye and Hiickel (I) who also found that, as the concen­
tration increases, the electrical fields of the ions inter­
fere increasingl y with each other. Consequently, local 
accumulations of a species of ions of varying concen­
tration will interact different internally and with sur-rounding reactants. The activity facton for ions in 
solution also vary by their different net electric charge 
in relation to the activity of surrounding media. In 
arldition, thermal factors, pressures, gravity and exter­
nal electric fields will, of course, also inlluence ionic 
dispersion. In a local region around any ion, the elec­
tric forces influence the movement of other ions. As 
concentration increases, the distance between ions de­
creases, which facilitates ion-ion interactions. The presented view of ionic potential has its corre­
spondence in the molecular potential of certain bio­
logically active nonionic compounds. Associated prob­
lems are particularly considered in the presentation of 
Ll-te ionar~rgonar concept of energy exchange over 
biologically closed electric circuits (Chapter XIII), an 
important prerequisite for the utilization of their bio­
logically stored electrical energy. This c. olldttued survey of some clraraeteris1ics of molecu­
lar and particulatt forces of attraction and repulsion may 
an jim imprmion umr far distant from problems of 
srrucrural rrwdif ~eation in biological materia. II is not so. These forces constitute importam portia/ functions in a 
complicattd systtm of many intmuring forces, all of which 
must be considertd in order to understand rek. ttd problems. Some of these functions will next be shown in stud­
i<S of movement of water in organized tissue (Chapter 
IX) and then of the behaviour of particles, i.
Short-range repulsive forces.
What type of forces are Vander Waals' forces?
The relationship is described by the function (p2/d3), where f is a function of angulation and d is the distance between the charges.
What is the relationship between the electrostatic forces and the distance between the particles?
The apparatus is used to study the electroosmotic water transport in tissue, and it consists of a chamber and two platinum electrodes.
What is the purpose of the apparatus in the electroosmotic transport experiment?
The experiment demonstrates the electroosmotic transport of water in human lung tissue.
What is the purpose of the experiment in Fig. IX: 3?
The relationship is not simply a matter of concentration, as the "total activity" of solute j is related to both concentration and other energy components, and the presence of net electrostatic forces on a molecule of water in the matrix largely depends on the spatial orientations and rotational moments of the particles.
What is the relationship between the concentration and the activity coefficient in the presence of charged particles?