One of the most important nonionic 
compounds, water, can be transported in a closed 
electric circut in a matrix lined with fixed charges. The 
mechanism of transport we can particularly refer to is 
evidently in the first hand represented by Type I 
(anomalous ) electroosmos is (Chapter IX). Dielectrics 
other than water may evidently also move in an electric 
field. This possibility will be demonstrated experimen­
tally in Chapter XIII. It is now suggested that the presence of vesicles in 
the endothelial cells indicates a mechanism of inter­
electrodic transfer of energy which is in addition to 
the mechanism of ionic transfer already proposed in 
Fig. XU: 30. Fig. XU: 3la shows the membranes of 
an endothelial cell and outlines !he suggested princi­
ple, that redox reactions lead to the development of 
basement membrane and fibrin ftlm as "external" pro-
INTERIOII Of CAPIUARY WALL 
BLOOD 
r.- LCotf>odle l!.l! )( )(~ 
.rf"" -"' '\ AnocUe Fig. XII: 31. (a) Suggested mechanism of energy 
transfer over capillary walls in addition to ionic 
transports illustrated in Fig. XII: 30. Endothelial 
cells are provided with electrode equivalent sites for 
reversible redox reactions at the outer and inner sur· 
faces of the endothelial cell membranes. As in any 
electrophoretic system, reaction products will de\•el· 
op at the electrode surfaces. It is !uggested that the 
basement membrane and the endothelial fibrin film 
have developed from reactions at the .. exterior" 
electrode surfaces. Vesicles are explained as non­
ionic collections of products of electrode reaclions, 
transported between the inner electrode surfaces, 
comparable to electrogenic transport of water in 
electroosmosis. (h) A vesicle produced at the inner 
surface of an endothelial cell is transported as an 
ergonar (see Chapter XIII) throuP, the cytoplasm, 
provided with matrix properties which permit inter­
electrode transport of dielectrics. On arrival at the 
opposite electrode. the nonionic comoound may ion· 
ize by a second electrode reaction. This pinocytotic 
.. pure" transport of water may be combined with hy· 
dration of, e.g., cationic compounds leading to en· 
hanced transpon of a variety of vesicular contents. R \... /-V..icular conecti«< of edox .. , .. non toni: compound 
mecti•tora Tr~,: • ~ (ergonlll") e.g. med••t , : 
enz~e m•trix : • moLul.,. , i =~~~tlonol 
,,.J·.,~Uonor) . . $_-_-.. ~ ~Cothod;o 
)(_(~) !_ 
e Anodic 
tNTI:RSTITIAL TISSUE 
Biologically closed electric circuits 145 ducts and vesicles as morphologically visible "inter­
nal" products in the VICC system. Electrode equiv­
alent redox sites at the endothelial cell membranes are 
anticipated to be reversible and therefore capable of 
allowing bidirectional ionic and vesicular (nonionic) 
transports across the endothelial cell. This mechanism 
aloo offers an alternative to the previously suggested 
Brownian type of vesicular motion. A panieular pat­
tern of distribution of vesicles adjacent to the inner 
walls of a cell (M, Fig. XII: 29b) can be regarded as 
representative of one phase of a unidirectional flow of 
current. One of the inner "electrode surfaces" should 
be anodic and the other cathodic. At a later stage when 
polarity and flow of current are reversed, a mixture 
will develop of vesicles. We have here assumed, for the 
sake of simplicity, that the vesicles arc transported as 
water in a closed electric circuit from anode to cath­
ode. A possibility might therefore be that vesi­
cles in pinocytosis are represented by matrix support­
ed transport of nticro-dustcrs of water molecules. Structural elements in the cytoplasm, such as fila­
ments, makes it at least justified to suggest that matrix 
elements exist to support the electrogenic transport of 
nonionic material. In this way a principle explaining 
vesicular transports now appears possible. Redox sines 
must be located on each side of endothelial mem­
branes, according to this theory for development of 
basement membrane, endothelial fibrin film and "e­
sjrular transport. These sites must each represent one 
*'exterior" and one "interior" reversible electrode 
equivalent surface (four redox sites). Such an arrange­
ment should allow bidirectional transport s of cations 
and anions as well as bidirectional transport of vesi­
cles. The direction of transport of a dielectric pack of 
energy (ergonar, see Chapter XIII) will be determined 
by the polarity of the surplus of fixed electric charges 
of the assumed matrix in the endothelial cytoplasm 
and the direction of the transcellular electric gradient. The principle of the development and unidirectional 
transport of a vesicle is presented in Fig. XII: 31 b. Several of the participating functions are still un­
known. The redox sites are assumed to consist of 
mediators such as enzyme molecules with the capacity 
to transfer electrons. Such models of electron conduct­
ing enzymes have also been described in the endotheli ­
al cytoplasm. The matrix properties of the cytoplasm 
are, however, still unknown in terrns of electrogenic 
transcellular transports. Related problems associated 
with transports of dielectric material should in princi­
ple be sintilar to the prerequisites for electrogenic 
transport of water (see Chapter IX). It should be 
evident that this model of vesicular transport also 
includes the mechanism represented by Type III elec­
troosmosis. Any cationic compound can be hydrated 
and transported in clusters as electropositive ergionars 
(see Chapter XIII). 146 Biologically closed electric circuits 8. Long and short distance selective transports in 
tissue over VICC systems 
To conclude this presentation, a survey of integrated 
events is presented in Fig. XII: 32. Metabolic processes or a local injury UJ a tissue will 
result in an electric (physicochentical ) potential differ­
ence in relation to the blood in surrounding vessels 
(Fig. XII: 32 a). The tissues then present their "de­
mand potential" for compensation of their internal 
metabolic imbalance. The electric potential difference 
delivers the driving electromotive force for closed cir­
cuit transports. This electromotive force consists of 
metabolic diffusion potentials (or diffusion potentials 
of catabolic reaction products from injured tissue) and 
redox potentials between blood and, e.g., hypoxic tis­
sue. Under the influence of the electric field, arterioles 
and arterial capillaries will contract in certain regions 
and possibly by means of the pericytes. Contraction of 
arterial capillaries may then lead to regional closure of 
leaking arterial stomata at the same time as corre­
sponding vdns are wide and continue to keep thdr 
stomata open. In Fig. XII: 32 b such selective arterio­
lar and arterio-capillary contractions are :seen in dog 
mesentery while venules and venous capillaries are 
wide and contain many leukocyte .s (Fig. XII: 32 c, 
mesentery electropositive, aorta electronegative, 0.1 
volt potential difference, 0.03 coulomb, haematoxylin­
eosin stain). A steep transcapillary gradient of an endogenous 
demand potential of a tissue in relation to blood should 
develop when the front of diffusion app:roachc:s the 
exterior of the capillary wall. Such a gradient is already 
present between the thrombus and blood (Fig. XII: 32 a). When the ntinimum potential difference 
across the capillary wall for the start of redox reactions 
is exceeded, electrode reactions will take place at the 
"electrodes" of the capillary walls. Each side of the 
endothelial membranes should have one anodic and 
one cathodic surface. Ion exchange will then start 
through the endothelial cell and over tlhe vascular 
interstitial channels of the VICC via leaking venules 
and venous capillaries. Packages (vesicles) of nonionic 
products (ergonars) from the endothelial electrode sur­
faces should be transported through the cyloplasm like 
water in electroosmosis (compare also pinocytosis ) and 
eventually become ionized ( = activated ) a.t the oppo­
site electrode. This prelintinary theory for long distance closed cir­
cuit transport between blood and tissue rep.resents only 
one of several possibilities. Evidently selective contrac­
tion of venules can take place also, although contrac­
tions in venules have been observed on[y rarely in 
these experiments. Selective closure of "enules and 
venous capillaries would probably also create rather 
extraordinary conditions, e.g., increase in capillary m-0 
Thrombus 
"---1+-f-Conduct ing 
tlnterstitiaf 
•fluid 
Dilated. leaking vessel 
-"' .. -c"" " ::.... ..... 
" .. \ 
~ • .. -
Fig. X/1:32. (a) Sch.ematic illustration ofVJCC in "micro­
injury" of tissue. Degrading products of an injured tissue 
produce diffusion potentials. The electric redox potential 
difference in relation to blood adds to the development of a 
field-induced regional contraction of adjacent arterial capil· 
!aries. The pores of arterial capillaries are then anticipated to 
close. Redox reactions will start at the outer and inner dec· 
ttode-equivalent surfaces of the endothelial cell membranes. Two layers of revers.ible redox sites are necessary at each cell 
membrane. Transpon of ions will start as long-distance 
transports over blood plastru~through open venous capillary 
pores, through interstitial fluid and as transports through the • "' ~ ' • ... 
~ • ' • • • . endothelial cytoplasm. Surfaces of thrombi may also serve as 
sites for redox reactions. (b) Simulation of the "injury poten· 
tial" difference in a, but without producing injury. One 
platinum electrode was gently positioned against the mesen­
tery of a dog and the other placed in a catheter in the aorta. 0.03 coulomb at 100 mV were passed between mesentery ( +) 
and aorta (-). Field-induced contraction of ar1erial capillar­
ies makes corresponding venules and venous capillaries fill 
by collaleral circulation of blood. The granulocytes (<, dark 
material in the wide venous branches) arc clectronegatively 
charged. They are attracted electrophoreti cally in the VJCC 
to the anode. Haematoxylin ·eosin. tration pressure and perhaps extensive diapedetic 
bleedings. The earlier suggestion that leaking, noncontracted 
capillaries should only allow minute closed circuits 
adjacent to the capillary membranes now can be recon· 
sidered and described as a mechanism for short-distan ce 
closed circuit transport across the capillary walls. These two systems of electrogenic transports appear 
to differ in principle by the mechanism of field· 
induced arteriolar and arterial-capillary contractions. Should this function be found in the future to be 
related to the presence of pericytes, the short-distance 
closed circuit transports should logically be localized 
preferably in regions where capillaries appear with 
only sparse numbers of pericytes (e.g., in the lungs). The present experiment s have also revealed a discre· 
pancy between morphologic descriptions and function 
of capillaries. Traditionally we distinguish anatomical· 
ly between arterioles , capillaries and venules. From 
the functional point of view it seems more logical to 
speak about leaking or contracting arterioles and arte· 
rial capillaries ("arteriocapillaries "). These capillaries 
are accompanied by predominantly leaking venous ca­
pillaires ("venocapillaries" ) and venules. Is there any further direct or indirect evidence that 
these anticipated functions of the capillaries exist? One 
argument might be that the clinical, radiologic and 
experimental observations reported in this book point 
to the existence of a VICC system. It seems difficult to 
find an alternative mechanism capable of explaining 
the multiplicity of biological events which can be ex· 
plaincd by the VICC system. Moreover, one may quts· 
rion if rissue function can "afford" nor 10 take advamage 
of rhe exceedingly efficient system for sekcrive transports 
which closed electric circuits offer. We will therefore 
continue in Chapter XIII with a presentation of how 
such circuits may be activated, which is closely related 
to the problem of transfer of energy. F. Conclusions 
Evidence bas been presented that biologically closed 
electric circuits (BCEC) ex.ist. These circuits appear to 
represent an additional , previously overlooked , circu­
latory system. One important specific circuit is called 
the VICC (vascular-interstitial closed circuit), which is 
parti~ularly considered in this study. This circuit is 
coupled to the mechanisms of bulk transport (diffu· 
sion and mechanical circulation ) as a system of selec­
tive transport. Beyond this capacity for selective trans­
port, BCEC systems are evidently able to influence 
"local" chemical processes in the circuits, taking place 
148 Biologically closed electric circuits also at "long" distances from the sources of the main 
driving forces. Thereby BCEC systems can be recog· 
nized to contribute to homeostasis. In later chapters 
other important functions of the interacting transport 
systems will be demonstrated, with particular empha· 
sis on structural development of tissue and healing 
processes. As an introduction to the description of the vascu· 
tar-interstitial closed circuit (VICC), two mechanism s 
have been distinguished for corrosion of metal im· 
plants in bone. The most common and well known 
mechanism of corrosion depends on locally different 
anoetic and cathodic parts of the implant, causing what 
is here called uncomplicated corrosion. The second 
mechanism is called complicated corrosion , in which 
relatively cathodic and anodic parts characterize tis· 
sues of different local electrochemical potentials adja­
cent to the metal. Levelling of these differences re· 
quires a closed electric circuit, which the metal partly 
creates. A closed circuit can, however, also be distin· 
guished in tissue without the presence of metal. Thts leads us to the concept of biologically closed 
electric circuits (BCEC). Several kinds of BCEC may 
yet be distinguished in the future, based on different 
structural arrangements of tissue components and of 
body fluids with different conductive properties. This 
chapter focuses on vascular-interstitial closed electric 
circuits (VICC). This type of BCEC has two conduct· 
ing main branches and an intersected regulating 
mechanism in the capillary walls. One branch is formed by the electrically insulating 
walls of "large" vessels surrounding their conductive 
component , the blood plasma. The other branch is 
formed by the conducting interstitial fluid and the 
insulating tissue matrix of cell membranes. The red 
blood cell membranes also possess a resistive function 
in the blood. These cell membranes therefore rcprc· 
sent a movable part of the matrix of the VICC, variable 
with the haematocrit. Electrical junctions between plasma and interstitial 
fluid are evidently present over the membranes of 
capillaries. An analysis of the capillaries with regard to their 
function in the VICC is of particular interest. The 
model for transfer of electrons between two redox sites 
of an enzyme molecule, suggested by Cope (10, II), 
has been tentatively accepted to explain the need for 
necessary electrode-equivalent redox sites of the 
VICC. The appearances of the basement membrane 
around the capillary and t/u endothelial fibrin film, 
facing the blood stream, ruggesrs rhar these srrucrures 
derive from endogenously droeloped products ar ouur 
"electrode" surfaces of redox sites located in the endo­
thelial membranes . The material of the basement 
membrane leaves an exoendothelial space open toward 
the endothelium. This space seems to correspond to the space ("A" zone) around a material under c:ontrac· 
tion by its own concentration forces. The underlying 
mechanisms are described previously in this chapter in 
connection with the analyses of in vivo corrosion in the 
presence of a tissue-matrix. The endothelial fibrin ftlm 
on the other hand appears different from the basement 
membrane, presumably partly due to the modifying 
effect of the streaming blood. These "external" prod· 
ucts of electrode reactions should have their corre· 
sponding "internal", intracellular products of reaction 
at the internal redox sites. The vesicles of the endothe· 
lial cells are suggested to contain such material. Ions 
tend to migrate separated in electrophoretic trans· 
ports. It is suggested that the vesicles, visible in elec­
tron micrographs, consist of mainly nonionic coUec· 
tions of electrode products. An electrogenic, closed 
circuit transport, even of pure nonionic material, is 
nevertheless possible when the transport is supported 
by a charged, intracellular "capillary" matrix. Struc· 
tural intracellular elements as fllaments possibly inher· 
it such functions. Intracellular transport of nonionic 
compounds (e.g., pinocytotic transport of water) may 
take place in a way analogous to the mechanism of 
Type I electroosmosis (p. 81). Such a mechanism 
should then represent an electrogenic , intracellular 
transport of energetic compounds in addition to ionic 
transports. Experimental support for the development 
and transport of vesicles in a matrix is also based on 
experimental analogues in Chapters IX, Xlii and 
XIV. Another model of vesicular transport is also 
suggested based on the formation of clusters ofhydrat· 
ed cations. The capillaries also seem to contain an interesting 
mechanism for switching between "short-dhtance" 
and "long-distance" VICC transport. When endothelial clefts (stomata) are open in the 
arterial capillaries, short-distance selective, ionic 
transports are possible over these leaking pores. Adja­
cent to their openings, the "outer electrodes " should 
provide electrode reactions and electron transfer to the 
"inner electrodes" of the membranou s redox sites and 
start ionic and vesicular transports through the cyto· 
plasm. By exposing mesentery of dogs to electric 
fields it was found that arterioles and arterial capillar· 
ies ("arteriocap illaries") contract and appear empty of 
blood cells in scattered vascular regions. Simulta· 
neously, corresponding venules and venous capillaries 
("venocapillaries ") fill with blood, evidently from 
collateral circulation from adjacent vascular areas 
without arteriocapillary contractions. The venocapil· 
!aries fill selectively with large numbers of granulo· 
cytes in dog mesentery near the mesenteric electrode, 
e.g., at 1-2 V potential difference . Granulocytes also 
pass by diapedesis through evidently open venocapil· 
lary pores to the interstitial tissue. These observations 
point to the creation of long-distance selective VICC transports of material when tissue is exposed to electric 
fields. It is suggested that the pericytes may be in· 
volved in the mechanism of regional closure of the 
pores between the endothelial cells. The electronegative granulocytes or an anionic com· 
pound, e.g., Evans blue dye, are electrophoretically 
attracted to the electropositive region of the tissue. Around an electronegative region in a closed circuit 
electronegative cells and compounds are repelled. In 
those veins which have their flow directed toward an 
electronegative focus, the repelling of granulocytes 
and anions in the blood stream causes them to accu· 
mulate in these veins. These observations contribute to 
the prc:scntation of an alternative explanation of so· 
called chemotaxis, which will be further treated in 
Chapters XIV and XVI. The behaviour of granulo· 
cytes in experimental activation of VICC channels il· 
lustrates the capacity of selective transports in tissue of 
this particular BCEC system. Ionization is a characteristic feature in tissue injury. For example, intravascular thrombosis is a type of 
injury which appears not to interrupt the conducting 
pathway. Indeed, intravascular conductivity is in· 
creased. The physicochemical potential of tissue injury 
represents a driving force, which over VICC channels 
leads 10 electrophoretic and electroosmotic transports 
and eventually to healing of the injury. Even small 
gradients of force in biologic material may be expected 
to produce significant changes if the "gates" are avail­
able, open and the time sufficiently long. Metabolic reactions not only form new products in 
tissue but also new electrochemical gradien ts in BCEC 
(VICC) channels. The products diffuse and migrate. They become dislocated in an electrochemically acti· 
vated closed circuit and then easily adsorbed at struc­
tural interfaces of tissue matrices in the circuit. In this 
way structures such as membranes may develop. The 
principle of this mechanism has been outlined for the 
development of intraabdominal organ capsules. The 
mechanisms of development of many other membrane 
structures , including basement membranes , arc little 
known but might be similarly explained. In future 
searches for analogues to electrodes in tissue, it seems 
likely that one of the places where they can be expect· 
ed to be found is adjacent to (fibrous) membranes . Biologically closed electric circuits (BCEC) is here 
used as a general term for structures with the capacity 
to channelize the exchange of energy in a selective way 
over long as well as short distances. In addition to the 
vascular-interstitial closed circuit (VICC) outlined in 
this study, other circuits can also be anticipated to 
exist, e.g., in conducting electrolytes in glandular 
ducts, cerebrospinal fluid, pleural and peritoneal fluid 
(with or without connection s tO vascular or interstitial 
conducting branches). The VICC system may be re· 
garded as a circulation for selective transport in tissue 
Biologically closed electric circuits 149 closely integrated with the nonselective bulk trans­
ports by diffusion and mechanical movement in the 
circulatory system. As a group of systems of selective 
transport, BCEC systems may be recognized as one of 
the pre~equisites for structural development and func­
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ionars and ergonars 
A. Components of BCEC 
systems 
Chemical reactions in vivo differ from chemical reac· 
tions in vitro by usually being isothermic . Biologic 
reactions are often modulated in many ways which are 
energetically economical and functionally highly dif· 
ferentiated. Acceptance of the principle of BCEC 
means that a variety of closed circuit functions, com· 
monplace in contemporary electronic technolog y, 
should also have their correspondence in biology. This 
correspondence includes not only the functions which 
drive the circuits hut also the influences of "locat'• 
reactions along electrochemical gradients over BCEC 
branches, either in series or in parallel. As will be 
seen, ionic and nonionic compounds each interact jn a 
way that makes selective distribution and modulation 
of energy possible over short and long distances. BCEC therefore represent, in the author's view, an 
important circulatory system in addition to the me· 
chanically driven circulation of blood. Components of 
this circulatory system are listed in Table XIII: I. Selective distributions of compounds thereby be· 
come possible, even against existing pressure gradi· 
ents, and may contribute, for example, to homeostasis. 152 Energetics , ionars and ergonars As far as can be seen now, the mechanical transport 
system and the electrochemical BCEC systems arc 
closely integrated both morphologically and function­
ally. Future evaluations of the function of the combi· 
nation of the two systems will, however, require a 
Table XIII: I. Componenrs of the circulawry system of BCEC 
I. Matrices Structured groupJ of channeli::i"t wmpontnll.' Vessels, duels, interstitial channels, fibrous mem­
branes. Some o( these components are contractile 
and change electrical rcsi$tan. :e . Cqnc~ rw1 «ggniuJ as clwrt~ls: 
Erythroc)'tes and other circuhaing ~us. celluJar 
c-omponents and ll\Oleeules. Ground substance, fi. brous material, nutrients, metabolites, metabolic 
wastes. Movable "matrices" add variable rcsis· 
lance and capacitance to the circuit.s. ll. Solnnt Water 
Ill. El«rrodr-cqwivalml inttr/aus for redox rtactUms 
IV. Ions I orr.an: 
CoiJtttlons of ions operating BCEC 
V. Ergons Erg(num: 
CoiJections of e.lecuicaJJy balanced tnergttk: mole· 
cuJC$ operating BCEC con&iderobly more differentiated knowledge of the 
BCEC systems than is presently available. The follow­
ing presentation will focus on the important problem 
of how BCEC systems can be activated and utilized in 
the transfer of energy in tissues. We will start with a 
general presentation of the author's view on the energy 
available for such functions. This introductory presen­
tation is intended only as a preliminary approach to­
ward understanding the associated complicated prob­
lems. These problems are indeed a consequence of the 
recognition of the concept of BCEC. The energy which can drive a BCEC needs first to 
be identified and defmed. One group of energetic 
compounds is primarily ionic and supplies electric (or 
more precisely, physicochemical ) energy to the sys­
tem. The other group consists of primarily nonionic 
energetic compounds. In order to facilitate handling of 
the actual problems it is now necessary tO introduce 
new, if tentative, terminology (Table XIII: 2). The term ergon (Greek, ergon = work) is used here 
to mean a nonionic compound possessing potential 
electric energy, as distinguished from an ionic com­
pound possessing immediately available electric ener­
gy, in a BCEC system. These two types of energy 
compounds are integrated in the energy conversion of 
BCEC systems as ionic and ergonic collections called 
ionars and ergonars, respectivel y. Together these col­
lections are termed ergionars. The need for a distinc­
tion between ionic and ergonic energies is a conse­
quence of the important differences of their energetic 
behaviour in a BCEC. Similarly, their appearance as 
ionars and ergonars has a particular meaning in closed 
circuit energy exchange. Examples of ergonars in­
clude oxygen, glucose, neutral fat, ATP, NADPH1 
and nonpolar amino acids, e.g., leucine, valine, methi­
onine and phenylalanine. Water may also be included 
in this group although it can be regarded in many 
respects as a special type of matrix material. BCEC systems require electric energy, supplied di­
rectly by ions or indirectly by ergons (after their acti­
vation). The electric potentials for redox half-reactions 
of ions and ergons can be defined by standard reduc­
tion potentials. In Table XIII: 3 some known redox 
potentials are presented (3, 20). Reduction or oxidation of ergons creates ions, which 
in turn by oxidation or reduction may cre-dte ergonic 
compounds. In this way we may regard ergonars as 
precursors of ionars and vice versa. Table XIII: 3 does 
not tell us when, where and under what biologic cir­
cumstances these reactions take place. The answers to 
these questions require further discussion of the con­
cepts of ionars, ergonars and BCEC. Ionars and ergonars represent two types of biologic 
bulk energy. Ergonars are electrically noncharged 
packages of energy. Their arrivals at places which are 
suitable for release of this energy depend mainly on Table XIII: 2. Energy in BCEC 1Y1tan.J: termmolog;y 
Unit Symbol 
hn (eltttricaUy 
charged moleeule) I!> 
Ergon {nonionic 
energy carrier) ~ 1-Energy C:l)llcctions 
lonar nx ~ I Ergjonar 
Ergonar ox~ 
diffusion and mechanical transport from the places of 
production. Glucose, for example, after liberation 
from glycogen in the liver, does not "waste" its energy 
during transport in the blood stream. On arriving at a 
suitable site of reaction, e.g., a working muscle, it 
contributes to the need of energy of the local tissue. During transport in bodily fluids, ions will create 
electromagnetic fields in surrounding tissue. Ergons 
do not. Selective electrophoretic transport of material 
according to electric gradients is the case with ionars. Such transport is not possible for ergonars. Neverthe ­
less, ergonars may undergo electrogenic transport, me­
diated by a suitable matrix. Differences of this kind 
appear to be important for differentiated transport of 
energttic compounds in tissue. An electrochemical 
gradient caused by ionars, e.g., from local metabolic 
activities in a tissue, should induce compensato ry, 
selective migrations of ionic material in a BCEC. If 
levelling of physicochemical gradients were to take 
place exclusively by mechanical transports (forced 
convection ), only an indiscriminate mixing should oc­
cur. Diffusion contributes to selective distribution. The tendency for selective distribution of ions in a 
BCEC also depends on the morphologic distribution of 
tile electrically inactive ergonars. The presence of er­
gonars adds a resistance and a capacitance in parallel in 
the circuits. The varying concentrations and locations 
of ergonars thereby modify the electric activities of the 
ionars. This mode of action is illustrated in Fig. XIII: I, where a part of a liquid-containing branch of a 
closed circuit (a) is conductive because of its contents 
of cations and anions and water molecules (crgons). Warer, funhermore, is a "nonionic" compound, 
which acts as a unique kind of movable matrix. At the 
same time it contains all the characteris tics of an er­
gonar of a complex nature. The capacity of water to 
form clusters of molecules of an energy content lower 
than the summation of energy of "individual" water 
molecules, emphasizes the importance of recognizing 
ergonars in energy conversion. The degree of breaking 
of clusters is influenced by many factors, e.g., the 
concentrations of ions, temperature and electric fields. The appearance of an ergonar other than the water, 
e.g., (b) a bolus of glucose, may markedly impair 
conductivity, while (c) the same quantity of ergonar in 
Energetics, ionars and ergonars !53 Table XIII: 3. Srandard redu.ri<m pormrials of'"""' oxidati<m-reducti<m luzlf-reactiom 
Re.ac-
rjon Half-reaction (wrinen as a reduction) 
L ~01+2H~+2c--+HtO 
2. Fe-''"+ le--Fc1-
3. 21"+2e--+l: 
4. Cylocbrome -o,-Fe'~ + le--
cytochrome -a)-Fe1 ~ 
S. S()! '+2W+2c· -SOJ'+H,O 
6. N0j+2H'+2c · -NOj+H10 
7. }02+H10+2c·-Ht01 
8. Cytochrome -a-Fe1' + te•--+ 
cyuxhrome-a-f'e1' 
9. Cytochrome -c-Fe*• + le -
cyuxhrorm-c-Fel' 
10. 2,6-0 iohlorophenolindopheno ~.,+ 
2H' +2e -2,6-0CPP,,.., 
II. Croronyi·S-CoA +lH' + 2< • -
buryryi-5-CoA 
12. Cu1' + le~-Cu .. 
13. .Melhemoglobin-Fe•· + Je--
he-moglobin- F~ · 
14. Ubiquinone+lH .. +l.e--ubiquinone-H2 
IS. Dehydroascorbate+2H ' +2e _. ascorbale 
16. Metmyoglo bin.-FeJ• + te·­
myoglobin-Fe1' 
17. Fumarau: +2H' +2c--succinate 
18. Methylene blue\ • .,+2H' +2c: --+ 
methylene blue(""• 
19. Pyruvate+NH1+2H'+2c· ~alanine 
20. a-Ketoglutarate+ NH3+2H. +2e--+ 
glutamate + H20 
21. Acetaldehyde+2H' +2e ~ethanol £0 at 
pH 7.0" 
(volts) 
0.816 
0.771 
O.SJ{i 
o.ss 
0.48 
0.42 
0.~0 
0.29 
0.2S 
O.ll 
0.19 
0.15 
0.1~9 
0.10 
0.06 
0.046 
0.0~0 
0.011 
-0.13 
-0.14 
-0.16~ Reac-
tion Half-reaction (written as a reduction) 
22. Oxalatttate+ 2H .. +2e--malate 
23. FAD+2H'+2e'-+FADH1 
24. Pyruvue+2H.+2e-~ lactate 
1S. Riboflavin + 2H" .. + 2e--+ ribo0avin-H1 
26. Cystine+ 2H .. +2e-~ 2 cysteine 
27. GSSG+2H' +2e· ->2 GSH 
28. S'+lH' +2e' -H,S 
29. l ,3-diph.osphoglya.ric acid+ 2H · + 2e--+ 
GAP+P, 
30. Acetoacetate+lH .. +le--
/1-bydroxybutyrar< 
31. Upoate,..:J+2H' +2e·-+ lipoate(n!IJ 
32a. NAO' +2H' +le'-NADH+H' 
32b. NAOP'+2H'+Z:e· -NADPH+H' 
33. Pyru\'ale+C01+2H.+2e--+malate 
34. Uric add+ 2H' + 2e--+ xanthine 
3S. Accryi·S-CoA+2H' +2c' .... 
""''aldehyde+CoA 
36. C01+lH' +2e·-+ formare 
37. n· + le-_. ~H1 
38. Ferredoxln-Fe '~ + te· -ferredoxin·Fe1• 
39. Gluconace +lH' +le--+ g:Jucose+HlO 
40. 3-Pbosphoglycc:nte +2H' +2e -
glyceraldehyde-3- ·phosphate + H10 
41. Meth ylvio logen~->+2H.+2e-­
methylvio loge~ ) 
42. Acetate+ZH .+2e--acetaldehyde 
43. Succinare+C01+2H'+2e -> 
a-ketoglutarate + H10 
44. Acetate+C01+2H.+2c -pyruvate ~at 
pH 7.0" 
(voll>) 
-0.17S -0. 18· 
-0.190 
-0.200 
-0.22 
-0.23 
-0.23 
-0.29 
-0.290 
-0.29 
-0.320 
-0.320 
-0.33 
-0.36 
-0.41 
-0.420 
-0.420 
-0.432 
-0.4S 
-o.ss 
-o.ss 
-0.60 
-0.67 
-0.70 
Source: I. H. Segel, Biochemical Calculations, 2d cd. (New York: Wiley, 197S), pp. 414-4lS. a Standard conditions: Unit activity of all component s except H*, which is maintained at 10-7M. Gases are at l arm pi'C$SUre. b The value siven is for free FAOIFAOH2. The £~of the protein-bound coenzyme varies. an incompletely obstructing location will impair con­
ductivity less. Conductivity also is affected (tf) by vari­
ations of morphology of the conducting vessels. In 
general, then, the functions of ionass and ergonars are 
variable, depending on their sizes and distributions in 
relation to the actual morphology of the BCEC 
branches. The intensity of closed circuit ionar interac­
tions now may be recognized to be modulated both by 
ergonars and by variations of the supporting noncon­
ducting walls around the conducting fluid. A purpose­
ful deceleration or acceleration of reactions over BCEC 
branches may be obtained in this way. (See also Fig. X: 6, as an experimental analogue of induced delay of 
spontaneous equilibrium ). Furthermore , any local nar­
rowing of a conducting branch of a BCEC should lead 
to local enhancement of current density, a change 
which may be expected to be accompanied by func-
I 54 Energetics, ionars and ergonars tiona! effects. A levelling of electric gradients over 
closed circuits between different regions of tissue 
should evidently also contribute to homeostasis. One way to determine the average balance between 
ionars and ergonars would be to determine their ratio 
of concentrat ion (elrgionar ratio) in the conducting 
medium. A more direct and simple way would be to 
determine the electric resistivity of the conducting 
medium. Complete analysis of ionar-ergonar interac­
tion in vivo would be extremely difficult to determine , 
partly for reasons mentioned above, i.e., functional 
variations of lumina of conducting branches as well as 
local convection factors. Moreover , one would need to 
know the morphology of associated, but different 
BCEC branches and systems as well as the vacying 
sites, magnitudes and times of occurrence of their 
energies of activation. a . -. + -+ . -.. -+ + 
+ • 
c 
-+ -
~ ~ Efoo-iia!i 
-~-·--'" . b 
d ~ .. --.. .. + -
+ + - -~ 
Fig. XIII: I. Interdependence of components of BCEC sys­
tems. (a) ElectricaUy conducting branch of a closed circuit, 
containing water and elec-trolytes. The vessel wall is electri· 
cally insulating. (b) Conductivity in this branch is markedly 
impaired by an ergonar (nonionic collection , e.g., a bolus of 
glucose) which locally fills the lumen. (c) Conductivit y is less 
impaired than in b because the same ergonar is more elon· 
gated and not occupying rhe entire cross seclion of the 
branch. (d) Impairment of conducti\•ity by compression or 
contraction of conducting closed c-ircuit branch. What are the mechanisms responsible for the dllfer­
ences of electric reactability of ergonars and ionars? This question can presently be answered only indirect­
ly by pointing toward some mechanisms which are 
likely to be involved. The magnitudes of ionic dissociation of different 
compounds and the buffering capacity of tissue fluids 
are evidently of importance for the electrical closed 
circuit events in a BCEC system. Depending on their different orbits of electrons, the 
elements differ in their ability to accept or donate 
electrons. Noble gases, which do not possess valence 
electrons, are therefore inert and unable to accept or 
donate electrons. Certain so-called noble metals, e.g., 
platinum, behave as inert materials by their property 
to form stable, electron-conducting surface films 
(PtO). Platinum is therefore often suitable as electrode 
material. The tendency for an element or molecule to 
deliver or to accept electrons in relation to another 
element or molecule is determined by what is called its 
electronegativity (17). The electronegativity of an ele­
ment is further characterized, according to Mulliken 
(12), by the difference between its ionizing energy and 
its electron affinity. These concepts are evidently of 
considerable interest with regard to the release of elec­
tric energy of ions and ergons. Ions carry an excess of electric charge immediate ly 
available for short and long distance reactions. Ergons 
do not "waste" their electric energy immediat ely and 
anywhere , e.g., during their transport. But at suitable 
sites of reaction they are able to release their energy 
when needed. Ergonars even cooperate along this line. Thus, glucose releases eighteen times as much energy 
in the presence of oxygen than in its absence (22). But oxygen, as an ergon, must protect its capacity for 
necessaty redox reactions during transport in the 
blood. This protection is possible partly as a. function 
of the microenvironment, which supports one of the 
ergonic characteristics (the saving of electric energy) of 
oxygen during its transport. The basic information 
which allows us to recognize this important mecha­
nism behind the enhanced ergonar function of oxygen 
is a result of extensive studies on the binding of oxygen 
to haemoglobin and myoglobin. A brief review of the 
relevant literature follows. The Structures of myoglobin and haemoglobin have 
been elucidated by Kendrew (8, 9) and Perntz (18). The capacity of myoglobin and haemoglobin to bind 
oxygen depends prin.cipally on a nonpolypeptide unit; 
the haem group. The haem is a prosthetic group of 
myo-and haemoglobin. It contains an organic proto­
porphyrin part and an iron atom, which can appear in 
ferrous (Fe++) or fe:rric (Fe•••) states of oxidation. Only ferrohaemoglobin can bind oxygen (29). Ferri­
haemoglobin is also called methaemoglobin. Corre­
sponding functions also apply to myoglobin (9). The 
prosthetic haem group in both myoglobin and hae­
moglobin is surrounded by the apoprotein, which in­
side consists almost entirely of nonpolar amino acids 
(27). Glutamine, asparagine, lysine and arginine, with­
out side chains, are present in the interior of the 
molecule (18). Residues that have both a polar and 
nonpolar part, such as threonine, tyrosine and trypto­
phan, are oriented so that their nonpolar portions 
point inward. Only 'two polar residues, each a mole­
cule of histidine, are present inside the myoglobin and 
haemoglobin molecule. Each of these histidines is very 
close to the iron in the haem group. The iron atom is 
directly bonded to one of the histidines (histidine F8, 
called the proximal histidine). The oxygen binding site 
of the iron is opposite the histidine F8, while the 
second histidine (E7, called the distal histidine) is 
nearby (22). In ferrimyoglobin the oxygen binding site 
is occupied by water, in deoxymyoglobin it is empty, 
in oxymyoglobin it i:s occupied by 02 (26). When 02 
occupies the binding site, it does not oxidize Fe•• to 
Fe+<•. How is this possible? The oxygen binding 
site of the iron atom is remarkabl y small in relation to 
the volume of the whole myoglobin molecule. This 
relationship gives a hint of the function of the polypep­
tide portion of the molecule surrounding the haem 
(25). The haem can be isolated. In a water solution it is 
able to bind oxygen ior a short time when its Fe • • is 
oxidized to Fe++•. Fe+++, however, can not bind 
oxygen. In some way the polypeptide poriion of 
haemo-and myoglobin stabilizes Fe • •. An answer to this problem of binding oxygen has 
been given by jai Wang (25) in an experimental mo­
del. When the haem group is imbedded in a polysty­
rene matrix containing a derivative of imidazole, it can 
Energetics, ionars and ergonars 155 undergo reversible oxygenation. The matrix prepara· 
tion mimics that of polypeptides. Wang proposes that 
imidazole has a role similar to that of the proximal F8 
histidine, while the polystyrene provides the haem 
group with a nonpolar hydrophobic environment. It is 
more difficult for electrons to leave ferrous iron in a 
nonpolar milieu than in water. The nonpolar haem­
binding site in myo-and haemoglobin therefore pro· 
tecrs ferrous iron from oxidation by exclusion of water 
(22). Myoglobin and haemoglobin are structurally simi· 
tar, but they are functionally different in their proper· 
ties as oxygen carrying molecules. For example, the 
affmity of haemoglobin for oxygen depends on pH, 
whlle that of myoglobin is independe nt of pH. In 
haemoglobin the oxygen binding is also affected by the 
affmity of haemoglobin for C02• Haemoglobin has a 
lower affinity for oxygen than myoglobin , an essential 
property from the point of view of transfer of oxygen 
from blood to muscle. When temperature or the con· 
centration of H .. or C02 increases, then oxygen is 
released from haemoglobin (Bohr effect), while these 
factors do not affect the capacity of myoglobin to bind 
oxygen (19). Several other factors are also involved in 
the mechanisms of binding and release of oxygen and 
C02 to haemoglobin . The point of this review may now be evident. The 
microenvironment of the haemoglob in molecule acts 
selectively to protect the ergonar oxygen from being 
ionized during its transport in the blood stream. Upon 
contact with myoglobin, the oxygen is transferred 
from haemoglobin and accumulated in the myoglobin , 
stiU in a protected , ergonic state until its oxidizing 
properties are needed. After thls introductory presentation of components 
of BCEC systems, it is now appropriate to consider the 
energy of ions and ergons. B. Ionic energy 
Current concepts of energy exchange in chemical reac­
tions will now be surveyed briefly, as a necessary 
prelude to considering biochemical reactions in the 
presence of biologically closed electric circuits. This 
survey is mainly based on the authoritative presenta· 
tions of Lehninger (10), Nobel (14), Bockris and Ora· 
l:ic (I), Morowitz (II) and Newman (13). The total energy·" of an ionic species j in a tissue is 
conventionally expressed as a sum involving its chemi· 
cal activity, volume and pressure conditions, electrical 
potential and gravity (14): 
Jt1= JtJ+RT1na1+V1P+z1FE+mjih (Eq. Xlll: 1). The potential energy I'J of the particular species j is 
156 Energetics, ionars and ergonars here related to a standard rderence level by the factor 
0 l'j. The next terrn represents chemical activity. R is the 
gas constant and T is temperature. The "effective 
concentration " is denoted as a1 of a species j, which 
will be experimentally examplilied in several sections 
of this book. a1 is a product of the activity coefficient 
Yi and the concentration c1. The chemical activity terrn 
RTina; is conventionally expressed in units of energy 
per mole. The effect of pressure-volume is expressed by the 
terrn V1P, where \11 represents the differential in· 
crease in partial molar volume after a differential 
amount of j is added. The effect of electrical potential is expressed by 
z1FE, where z1 represents the charge number ofj, F is 
Faraday's constant and E the actual electrical poten· 
tial. The last term m~h includes an expression for grav· 
ity, where g is the gravitational acceleration factor and 
h the height of a mass m in relation to a reference level. In ealculations and descriptions of biologic energe· 
tics, Gibbs' free ene.rgy (G) expresses the maximum 
sum of different energy qualities whlch are available 
for work. Gibbs' free energy of a system therefore 
consists of many components, whlch with regard to 
ionic energy in a tissue may be expressed as: 
G = 2: njl'j (Eq. Xlll: 2). i 
n1 represents here the number of moles of species j in 
the system and I'; the physicochemical potential of 
species j. These components of stored energy may be 
released either by individual chemical reactions be­
tween two reactants or by a series of reactions. In 
simplest form, A and B may react, producing C and 
D: 
aA+bB ~cC+dD 
wher< a, b, c, d represent participatory moles of A, B, 
C, D. 
The bulk energy change of the reaction expressed by 
t.G is then expressed: 
(Eq. Xlll: 3). The signs of this equation are determined by a de· 
crease (negative) of reactantS and an increase (positive) 
of the products. t.G represents in Eq. 3 the bulk 
change of free energy, whlch, by considering the indi· 
vidual energy factors of Eq. I can be expressed 
t.G =-af.l~ -bp~+Cjt~+d}l~ 
+RT( -a In a A-bIn a8+c lnac+dln a0) 
+P( -av.-b"8+d c+dV 0) 
+FE( -azA -bz8+czc+dzD) 
(Eq. Xlll: 4). Eq. 4 is somewhat troublesome to handle in practice. Simplified versions are therefore sometimes accepted. Most chemical tissue reactions, for example, take place 
at constant temperature and often at constant pres­
sure. If the volume of the reactants (all11+bV8) is the 
same as the volume of the products {eli 11 +dV 0) the 
pressure-volume expression in Eq. XIII: 4 is :rero. Moreover, the sum of the terms in the second to last 
parenthesis in Eq. XIII: 4 equals zero because no 
charge is created or destroyed, i.e., the term 
(az11+bz8) equals the term (czc+de0). The last par­
tial reaction in Eq. XII I: 4 also is zero because no mass 
is created or destroyed by the reaction. The constant 
terms of Eq. XIII: 4 (-a,u~-b;t~+CJt~+dp~ can be 
replaced by tj,Cf'. Eq. XU! : 4 can now be simplified 
to: 
(Eq. Xlll: 5). When local chemical reactions are of the type pre­
sented in Eq. XIII: 5, their electrical energy is convert­
ed berween the reactants at short distances of contact. In an electrolyte solution separated ions of opposite 
charge balance each other electrically. In spite of the 
absence of net electric charge the ionic separation is 
one of the prerequisites for new ionic recombinations. For local chemical reactions, the electrochemical po­
tential is often expressed in the simplified version of 
Eq. XIII: I as follows: 
(Eq. XIII: 6). Formulas for calculation of electrochemical energy in 
the transfer of ions include oxidation- reduction reac­
tions, where the redox potential E; may be written: 
£.=£?-RT In (reduced}) 
' ' zF (oxidized) ) {Eq. XIII: 7). C. Interdependence of 
energies, including gravity 
The abbreviations leading to Eq. XIII: S-7, useful for 
chemical in vitro reactions, may be regarded from a 
slightly different point of view in in vivo situations. It is easy to recognize that oil four partial energy 
factors of Jli (Eq. XIII: I) are interdependent. For 
instance, a local change in concentration of ions will 
influence local volume-pressure conditions, and vice 
versa. The piezoelectric properties of dry and living 
bone tissue probably reflect such connections . Exam­
ples from physiologic studies of membranes are simi­
lar. Such considerations led Teorell (23, 24) to de­
scribe the i.ntegrated functions of chemical energy, 
12-824586 Nord...s-volume-pressure and electric energy of a system he 
called the membrane oscillator. This concept repre­
sents an excitable system which is able to convert 
electrochemical energy into mechanical work. As in­
terdependence of all four energy factors, and panicu­
larly of gravity, is not immedia.tely obvious, the factor 
of gravity will now be the subject for some consider­
ation. It may seem unconventional to include a gravitation ­
al term in Eq. XIII: I. Gravity is, however, an impor· 
tant and of!en overlooked factor. Small but constant 
forces are known empirically to produce considerable 
effects in vivo when they are allowed to act over long 
periods of time. Thus, the relative distribution of air 
and blood in lungs depends, for example, to a large 
extent on gravity. Gravity is known to affect the ratio 
between ventilation and perfusion in upper and lower 
parts of the lungs (28). Even the ratio between system­
ic and pulmonary arterial flow can be anticipated to 
vary in different parts of the lungs as a function of 
gravity. Gravitational forces are relatively less influen­
tial on blood in the high pressure bronchial arteries 
than in the low pressure pulmonary arteries. Weight of 
the blood and, to some extent, of the lungs themselves 
makes the peripheral air spaces larger at the apices of 
human lungs than at the bases. Gravity-induced rela­
tive effects on ventilation and on the pulmonary and 
bronchial circulations in different parts of the lungs 
can therefore be expected tq trigger biological mecha­
nisms which are of importance for the physiologic and 
pathophysiologic distribution of blood flow and venti­
lation. For example, well-venuilated portions of lung 
trigger their divisions of the pulmonary arteries to 
dilate and preferentially distribute blood to them (5). A shunting of systemic anerial blood, as from the 
bronchial arteries, to precapillary pulmonary vessels, 
will, however, constrict local pulmonary vessels. Gravity has long been known to influence the local­
ization of pulmonary tuberculosis. Thus, pulmonary 
tuberculosis in man is found most commonly in the 
apices of the lungs, while in the bat, which hangs 
upside down during hibernation, the disease tends to 
occur in lung near the diaphragm. In the cow, pulmo­
nary tuberculosis is preferentially dorsal. When the left side of the bean fails to pump ade· 
quately, the resultant oedema distributes preferentially 
to dependent parts of the lung (and other dependent 
parts of the body), a well known reaction related to 
gravity. Orthostatic reactions, balance and orientation 
are other simple and familiar examples of gravitational 
forces in animals. Deep diving animals such as whales are exposed to 
very high external pressure when they dive as deep as 
70 m under the surface of the sea. The external water 
pressure then compresses peripheral capillaries, pro­
moting central diversion of blood. To prevent overload 
Energetics, ionars and ergonars I 57 on the hear! and lungs, these animals possess valves 
which regulate the amount of venous blood that is 
allowed to flow into the thora.x. aud l'ea.ch the bean.. It 
is in fact easy to fmd many morphological and func· 
tional adjustments to the force of gravity in animals as 
well as in plants. Furthermo re, the force of gravity can 
be utilized in angiography to modify flow in vessels 
and to direct contrast material into predetermined 
vascular regions (IS, 16). The author is awaie that 
most gravitational effects of the type described above 
may be considered as "systemic", meaning that the 
mass of any individual molecule is so small that the 
gravitational force is apparent only on large, concen· 
trated collections of molecules of different masses. However, this view could then also be applied to the 
chemical potential whose activity coefficient is depen· 
dent on concentration. For example, the gravitational 
effect of external sea water on distribution of blood in 
a whale, 70 m deep in the sea, should not be regaided 
as simply an effect of the weight of water. It is the 
gradient between reactants that counts in any concept 
of potential. The main message of this discussion on gravity is 
that this factor must not be neglected in considering 
the truely "systemic" BCEC reactions. VICC, e.g., 
represents an addi!Umal electrogenic circulatory system in· 
regrated with the mechanical bulk transport system of 
blood and lymph circulation . Ionars and ergonBIS may 
present small but important volume-pressure and 
gravitational forces. The effects of even small gradi· 
ents in vivo are partieulaiiy influenced by the factor of 
time. The "electrochemical" simplified Equation 
XIII: 6 is practical and useful in most in vitro reac­
tions. In biology, when reactions in vivo take place in 
highly specialized matrices and over long time periods, 
aU four energy components of the total physicochemi· 
cal potential must be considered. The term "electro­
chemical" is, however, often used in the broad sense 
of "physicochemical" when electric and chemical en· 
ergies are dominating. D. Ergonic energy 
The principle of BCEC makes it neceswy also to 
consider the interactions of noncharged species as an 
integrated factor in closed circuit functions. These 
interactions lead us to the following considerations: 
Whereas an ion is electrically charged, the electrical 
state of an ergon is internally balanced but electrically 
unstable. Ergons, like ions, are carriers of physicoche· 
mical energy. The total energy of an ergonic species e 
might then be expressed: 
1', = .tt~+RTlna , +[z,.FEJ*+ V,P+m,gh (Eq. XIII: 8). The total ergonic energy has the electric component 
I 58 Energetics, ionars and ergonars = [z,.FE)*.
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:

        One of the most important nonionic 
compounds, water, can be transported in a closed 
electric circut in a matrix lined with fixed charges. The 
mechanism of transport we can particularly refer to is 
evidently in the first hand represented by Type I 
(anomalous ) electroosmos is (Chapter IX). Dielectrics 
other than water may evidently also move in an electric 
field. This possibility will be demonstrated experimen­
tally in Chapter XIII. It is now suggested that the presence of vesicles in 
the endothelial cells indicates a mechanism of inter­
electrodic transfer of energy which is in addition to 
the mechanism of ionic transfer already proposed in 
Fig. XU: 30. Fig. XU: 3la shows the membranes of 
an endothelial cell and outlines. he suggested princi­
ple, that redox reactions lead to the development of 
basement membrane and fibrin ftlm as "external" pro-
INTERIOII Of CAPIUARY WALL 
BLOOD 
r. - LCotf>odle l. l. )( )(~. rf"" -"' '\ AnocUe Fig. XII: 31. (a) Suggested mechanism of energy 
transfer over capillary walls in addition to ionic 
transports illustrated in Fig. XII: 30. Endothelial 
cells are provided with electrode equivalent sites for 
reversible redox reactions at the outer and inner sur· 
faces of the endothelial cell membranes. As in any 
electrophoretic system, reaction products will de\•el· 
op at the electrode surfaces. It is. uggested that the 
basement membrane and the endothelial fibrin film 
have developed from reactions at the. exterior" 
electrode surfaces. Vesicles are explained as non­
ionic collections of products of electrode reaclions, 
transported between the inner electrode surfaces, 
comparable to electrogenic transport of water in 
electroosmosis. (h) A vesicle produced at the inner 
surface of an endothelial cell is transported as an 
ergonar (see Chapter XIII) throuP, the cytoplasm, 
provided with matrix properties which permit inter­
electrode transport of dielectrics. On arrival at the 
opposite electrode. the nonionic comoound may ion· 
ize by a second electrode reaction. This pinocytotic. pure" transport of water may be combined with hy· 
dration of, e. g. , cationic compounds leading to en· 
hanced transpon of a variety of vesicular contents. R \. /-V. icular conecti«< of edox. ,. non toni: compound 
mecti•tora Tr~,: • ~ (ergonlll") e.
It is suggested that the presence of vesicles in the endothelial cells indicates a mechanism of interelectrodic transfer of energy, which is in addition to the mechanism of ionic transfer already proposed in Fig. XU: 30.
What is the mechanism of interelectrodic transfer of energy in the VICC system?
The endothelial cells are provided with electrode equivalent sites for reversible redox reactions at the outer and inner surfaces.
What is the purpose of the endothelial cells in the VICC system?
Ions carry an excess of electric charge immediately available for short and long distance reactions, whereas ergons do not "waste" their electric energy immediately and can only release it at suitable sites of reaction.
What is the difference between ionic and ergonic energy?
Gravity is an important and often overlooked factor that can influence the distribution of blood flow and the development of certain biological processes.
What is the role of gravity in the BCEC system?
Water is a non-ionic compound that can be transported in a matrix and is used as a matrix material to support the electrogenic transport of nonionic material.
What is the significance of the presence of water in the BCEC system?