Life at the Cell and Below-Cell Level. The Hidden History of a Fundamental Revolution in Biology
Gilbert N. Ling, Ph.D.
Pacific Press
ISBN 0-9707322-0-1

"Dr. Ling is one of the most inventive biochemist I have ever met."
Prof. Albert Szent-Györgyi,
Nobel Laureate

Chapter 8.

Aftermath of the Rout
(p. 40-42)

The long-term outcome of this rout of the protoplasm-oriented cell physiologists by A. V. Hill was devastating. The iconoclastic and often brilliant ideas and courageous voices of Martin Fischer, Herbert Roaf, Benjamin Moore, Ross Gortner, W.W. Lepeschkin vanished from the scene. It is in this "purified" silence that my career as a cell physiologist began in Chicago.

Only many years after receiving my Ph.D. degree did I discover piece by piece that there were alternatives to the membrane theory. And still later, did I realize how virtually all protoplasm-oriented cell physiologists in America and Europe had raised their heads brightly for a moment and then vanished. That is, with the exception of two isolated groups in far away Communist eastern Europe. One was led by E. Ernst in Pécs, Hungary; the other by Dmitrii Nasonov in Leningrad of the former Soviet Union.

8.1 The tiny Hungarian enclave under E. Ernst

Like Fischer, Moore, Roaf and others before him, Eugene Ernst (Ernst Jeno) (1895-1981) also believed that the membrane theory was wrong.83 Nor did he believe that cell water is normal liquid water and that cell K+ is free.

Instead, he believed that cell K+ is largely unionized, undissociated and unhydrated83 p 152 Ernst subscribed to the view of Macallum and others [10.2(5)] that K+ in striated muscle cells is not evenly distributed but located in the A-bands.84 And that a substantial part of cell water is "swelling water" (Schwellungswasser, Quellungswasser) associated with muscle cell proteins.399 In support, Ernst and his coworkers showed that both
NaCl and gelatin lower the relative vapor pressure of water, but gelatin does it many times more effectively than NaCl. From this perspective, they argued that frog muscle more closely resembles gelatin-gel than a solution of NaCl.83 p 112; 399

In anticipation of work to be described below [11.2], I may mention that Ernst's study of water sorption of frog muscle and gelatin was at a much lower range of relative vapor pressure than that of the normal physiological environment. In fact, most of their data points on frog muscle correspond to a final water content 10% or less of the total water content. The few data points corresponding to higher vapor pressures are too scattered to assess their true significance. This scattering could reflect too short an equilibration time (2 to 3 days),399 (see end of [11.2]).

Professor Ernst died in 1981. His work was continued by his student, Professor Joseph Tigyi and later by Professor Miklós Kellermayer and his group at the University of Pécs.85 In 1994, the University of Pécs kindly awarded me an honorary doctoral degree.

8.2. The Leningrad school under Nasonov and later, Troshin.

Dmitrii Nasonov (1895-1957) was born in Warsaw, the son of a zoology professor. Nasonov started out as a microanatomist, studying at one time at Columbia University in New York under cytologist E. B. Wilson, mentioned earlier. Nasonov was a decorated war hero in the siege of Leningrad during World War II. After the war, he returned to science and held the position of the Director of the Institute of Cytology in the same city.

With a solid foundation of cell anatomy, Nasonov's career as a cell physiologist was distinguished by his unwavering conviction that a sound cell physiology must rest upon a correct cell anatomy. And that a correct cell anatomy is impossible without taking into account that the cell is solid and made of protoplasm. His protein theory of cell damage and excitation illustrates the general direction of his own and his immediate associates research.86

Nasonov also introduced his phase theory of permeability and bioelectrical potentialscontending that cells do not possess a cell membrane with varying permeability86 164 and that an electrical potential difference occurs only across the surface of injured protoplasm but not across the surface of normal resting cells. This is how Nasonov described it in his own words: "According to our theory, the electromotive force arises only at the moment of injury or excitation when the electrodes (electrolytes ?, query of G.L.) are released from their linkage with the protein substrate. In this respect our theory resembles the alteration theory of Hermann (1885)." 86 p 178; 87; 15 p 20

I disagree to varying degrees with Nasonov on these two specific concepts. Thus the two-fold increase of permeability to sucrose following cell amputation offers one piece of evidence for the existence of a diffusion barrier (or cell membrane) at the normal cell surface (Table 1, see also [13.6]). The exploration with the Gerard-Graham-Ling capillary glass microelectrode88 makes it hard to deny the existence of a potential difference across the healthy normal resting cell membranein harmony with the "pre-existence theory" of cell potentials first offered by L. Hermann's teacher, Emil DuBois-Reymond.89 I shall return to the question of why Nasonov believed that there is no cell membrane.

Differences in some specific issues notwithstanding, I have great admiration for the courage and originality of this remarkable investigator. His, Aizenberg's and Kamnev's demonstration that permeant solutes can cause sustained shrinkage of living cells described in [4.1(3)] is but one of their landmark discoveries. Other major contributions from the Soviet school of cell physiology are described in his monograph, "Local Reaction of Protoplasm and Gradual Excitation"86 and in his student, A.S. Troshin's monograph, which will be reviewed next.

The Institute of Cytology at Leningrad survived Nasonov's death in 1957 and its directorship went to his prize pupil, Aphanasij S. Troshin, or simply A.S. Troshin.


"Life at the Cell and Below-Cell Level.
The Hidden History of a Fundamental Revolution in Biology":

Contents (PDF 218 Kb)
Preface (
PDF 155 Kb)
Answers to Reader's Queries (Read First!) (
PDF 120 Kb)

1. How It Began on the Wrong Foot---Perhaps Inescapably
2. The Same Mistake Repeated in Cell Physiology
3. How the Membrane Theory Began
4. Evidence for a Cell Membrane Covering All Living Cells
5. Evidence for the Cell Content as a Dilute Solution
6. Colloid, the Brain Child of a Chemist
7. Legacy of the Nearly Forgotten Pioneers
8. Aftermath of the Rout
9. Troshin's Sorption Theory for Solute Distribution
10. Ling's Fixed Charge Hypothesis (LFCH)
11. The Polarized Multilayer Theory of Cell Water
12. The Membrane-Pump Theory and Grave Contradictions
13. The Physico-chemical Makeup of the Cell Membrane
14. The Living State: Electronic Mechanisms for its Maintenance and Control
15. Physiological Activities: Electronic Mechanisms and Their Control by ATP, Drugs, Hormones and Other Cardinal Adsorbents
16. Summary Plus
17. Epilogue 

A Super-Glossary
List of Abbreviations
List of Figures, Tables and Equations
References (
PDF 193 Kb)
Subject Index
About the Author

"Life at the Cell and Below-Cell Level..."
"Gilbert Ling"