R L Maynard
Honorary Professor, Birmingham University, UK

https://doi.org/10.36866/pn.94.9

The age of great textbooks of physiology seems to have passed. Those splendid thousand-page volumes that used to inform and perhaps intimidate have, like battleships, disappeared. But anyone who studied physiology 40 years ago might regret their passing and those who are now studying the subject are deprived of a resource and, perhaps, a pleasure. I remember the step change from introductory textbooks for medical students to books for students of physiology per se: Davson’s two volumes on General Physiology, Ruch and Patton, Mountcastle’s two large volumes, Davson and Eggleton’s 14th edition of Starling’s Principles of Human Physiology, Bayliss’s new edition in two volumes of his father’s General Physiology and Best and Taylor’s 1800 pages. Not one of these works remains in print but in an age when ‘reading physiology’ meant reading physiology they were well known. Of course our subject has moved on and these books are now regarded as out of date. This is not entirely true: much may still be learnt from their pages. In this series I shall discuss just a few of the ‘greats’ from 40 years ago.

Ruch and Patton (Physiology and Biophysics, 19th edition, published by W B Saunders in 1965) was a superb book: perhaps the best of the textbooks of the 1960-1975 period, perhaps the best of the 20th century. The 19th edition was dedicated to John F Fulton who had taken over the 15th edition of what had been in 1896, An American Textbook of Physiology edited by W H Howell. Fulton was joined by T C Ruch for the 18th edition and the word ‘Biophysics’ was added to the title. Two further editions, the 20th (in three volumes) and the 21st were produced and then it died. Copies of the great 19th are still available at very modest prices and are worth hunting out. What made it so good?

The answer was and is obvious: the distinction of the authors and the quality of their writing. J Walter Woodbury, his son Dixon M Woodbury, Harold Copp, TC Ruch, HD Patton, RF Rushmer, RL Riley, AM Scher: a roll call of leading physiologists all at the top of their form. The book was not an easy read: I remember reading in the Preface, “This is not an easy textbook for students, nor is physiology an easy subject lending itself to memorization. Students… may find it initially difficult, but with application, the know-how comes and with it the pleasures of following the quantitative pathway… ”. Space forbids a lengthy review but two contributions are worth a little discussion.

J Walter Woodbury’s chapters on electrophysiology (membrane potential and action potential) were outstanding and still read very well today. The author set out to explain, with quantitative rigour, what Hodgkin and Huxley had discovered: the excitement was tangible. Woodbury had been influential in recommending Hodgkin and Huxley for the Nobel Prize and understood their work so very well. His writing sparkles with intelligence and wit. An appendix “for use by readers whose physics is rusty” cost me a lot of paper and pencils: the problems are worth looking at today. An easy read? Hardly that, but the quality of the explanations was memorable: Figures 7, page 21, and 4, page 35, deserve to be embossed on the walls of physiology departments. And it was witty! There was no sign of the patronising approach one found in other books; everything was explained, step by step, with numbers and equations. Anybody who wishes to find a place to start in understanding electrophysiology and who is foxed by the difference between a capacitance current and an ionic current would be well advised to read Woodbury. His chapter ‘Regulation of pH’ was just as good: if one word sums it up that word is rigour. But the rigour again tempered by wit: pH? logarithms? Calculate your own! All you need to remember is 0.3010, 0.4771 and 0.8450 and you can work out the rest in your head!

Riley’s chapter, ‘Gas Exchange and Transportation’, was equally good. Riley, Rahn, Otis, Fenn, Fahri and Cournand were the great men of respiratory physiology in the 1950s. In his chapter, Riley began at square one and wrote out a summary of work on gas exchange. A summary? Yes, the full account appeared in the great American Handbook of Physiology, but what a summary! The chapter bristled with equations, properly developed and derived, and with graphs that actually explained things. The development of the CO2-O2 diagram was so well explained that I remember the thrill I experienced when I realised, for the first time, how to combine the gas and blood R lines.

How much of this is worth reading today? I think a lot of it is worth reading: it provides a starting point for understanding what has happened since but, and more important, it provides such a good example of what physiology is actually about. Physiology is the explanation of biological phenomena in the terms of physics and physical chemistry. A quantitative approach is essential. Ruch and Patton took that approach and their book remains a landmark in the history of textbooks of physiology.

Editorial note by David Miller: The sentence in Bob’s last paragraph; “Physiology is the explanation of biological phenomena in the terms of physics and physical chemistry” is very close to Davson’s definition of general physiology as; “the study of those aspects of living material that show some immediate prospect of being described in the terms of the known laws of physics and chemistry”. It is fascinating that this sentiment chimes well with the views of Sydney Ringer, no less. When he chaired the Society Dinner at the March 1891 meeting, a motion was passed stating ‘’That in the opinion of this Society it is important that, as recommended by the General Medical Council, a large part if not the whole of the additional year which is to be added to the medical curriculum should be devoted to Elementary Physics, Chemistry, and Biology.”