Tuesday 11 June 2024

The Heart of the Salamander: Science in the Making 1940-41

The Royal Society sometimes publishes papers from the archives under the heading Science in the Making. A recent one certainly drew my attention since it is a letter from 1940 concerning a paper and people I have written about previously. It provides an amusing insight into how papers were refereed and of why the whole business of selecting scientific papers for publication by peer review is so fraught with difficulty, why mistakes are often made and how good papers are sometimes rejected while utter garbage gets published sometimes, sadly, by what were/are highly-regarded journals. Before I digress into a tirade on the decline in standards of present-day refereeing and editing, back to the paper in hand.

The letter from the archives was written from Trinity College, Cambridge on 28 December 1940 by Carl Frederick Abel Pantin FRS (1899 –1967) of the Department of Zoology. From the letter’s content it is obvious that he had been asked by John David Griffith Davies (1899-1953, the Assistant Secretary from 1937 until 1946) to provide a second opinion on a paper submitted for publication. Griffith Davies was trying to keep the publication system working as war took its toll and he was running from pillar to post moving records to safety, compiling a register of scientists for the war effort and running the London operation. The first refereewas Alan Nigel Drury FRS (later Sir Alan) (1889-1980) who recommended publication but that the introduction should be shortened. Here I declare a personal interest since after Sir Alan retired for the first time from the Lister Institute and having established a blood transfusion and blood products system during the war, he was persuaded to become head of the pathology department at the then new Institute of Animal Physiology at Babraham. After his second retirement he still kept a close interest in the people and work of the Institute. Although he was then 88 and frail, on my last day at Babraham before leaving for Ayr he asked to be picked up from home and brought to my lab in order to wish me well.

The paper in question was The Heart of the Salamander (Salamandra salamandra, L.), with Special Reference to the Conducting (Connecting) System and its Bearing on the Phylogeny of the Conducting Systems of Mammalian and Avian Hearts by Francis Davies (1897-1965) and Eric Thomas Brazil Francis (1900-1993) of the Departments of Anatomy and Zoology, respectively of the University of Sheffield.



Francis Davies
Eric Francis












Pantin’s first criticism was that the historical introduction was far too long. ETBF was following the tradition set by his former professor of zoology at Reading, Francis Joseph Cole FRS (1872-1959) with regard to immensely long histories of the subject of the publication. Pantin and Drury were right.

On the second point, that the paper needed to refer the paper to somebody from a human anatomy department, since the work would only be of interest to anatomists, was wrong. This anatomical paper, with as much physiology as the authors could do at the time, was about a physiological and evolutionary question. In short, as the title states it was about the conducting system within the heart responsible for the co-ordinated contraction of the heart muscle. In mammals and birds there are specialised muscular fibres within cardiac muscle which conduct the signal to contract from the pacemaker in the sino-atrial node to all parts of the ventricle. However, Davies and Francis could found no trace of these tracts, like the Bundle of His, or the smaller Purkinje fibres, in mammals. Thus they concluded that the wave of contraction that passes through the amphibian heart is the result of one contracting cardiac muscle cell setting off its neighbour, as in a line of falling dominos.

The implications for the way the heart works has very great implications in terms of the force of contraction and maximum heart rate that can be achieved. The wave of contraction is slow in amphibians compared with the rapid conduction to all parts of the heart in mammals and birds. In turn, the rate of metabolism of amphibians and reptiles is constrained making them ‘slow animals’ as opposed to the mammals and birds which have higher metabolic rates. There has been some suggestion that the development of specialised conducting fibres may also be related to the complete separation of the two sides of the heart in mammals and birds since in crocodilians, which have a full septum between the ventricles, there appears to be the rudiments of a conducting system. Davies and Francis later turned their attention to the crocodilian conducting system but that is a story for another time.

Despite Pantin missing the point completely, the paper was then sent to a third referee of the background Pantin suggested,  James Thomas Wilson FRS (1861-1945), Emeritus Professor of Anatomy in Cambridge. Although he was ill and felt unable to provide a detailed report, he considered it ‘a work of considerable interest and importance’. Wilson was a good choice. He had a reputation as a comparative anatomist with considerable knowledge of monotremes and the evolution of mammals gained while working at the University of Sydney.

The Davies and Francis paper was published in Philosophical Transactions, with a historical introduction that was still too long. It, together with later work on crocodilians, is still widely quoted since Davies and Francis were onto something of major importance in the evolution of vertebrates (Pantin was very much and invertebrates man) and of how animals work and why they work the way they do. The working of the heart, sadly only possible to address by exercises in armchair physiology, is also vital to understanding how dinosaurs managed to work at all.

The letters and reports from Drury, Pantin and Wilson certainly provide a snapshot from 1940 of science in the making.


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