How Chameleons Work: Great discoveries in the 1930s by Alexander Zoond aka Alexander or Alec Sand (1901-1945)

SPLAT. KERPOW. The words of a schoolboy comic hardly do justice to the chameleon launching its tongue to capture its prey. That whole process, from moving the eyes into stereoscopic mode, preparing the tongue for launch, the explosion of the tongue from the mouth, to the prey being dragged into the mouth and chewed must surely rank as one of the great sights in the living world. Equally the colour-changing abilities of these lizards has to be seen in real time to be appreciated.

Sadly, and the story even ends on a very sad note, the name of the man who made the most important discoveries in how the chameleon tongue works and of how colour changes are controlled has been largely forgotten. Partly, this is because he changed his name and partly it is because he died young and in tragic circumstances.

The name of the author that appears in list of references to the work on chameleons in the early 1930s is A. Zoond. The only reason I knew that he changed his name is that ETB Francis told me so; he did that while we were talking about his work on chameleons I had supplied him with. Zoond, he said, changed his name to Sand. In fact both names appeared on the extensive reading list ETBF handed out to honours zoology students for his course on vertebrates, a list so extensive a modern student would have an apoplexy, a. from shock over the quantity, and b., from having to work out whole fields of research for themselves.

Alexander Sand
In the tropical, white, uniform of a
Lieutenant, R.N.V.R.
Photograph used in his Royal Society
Obituary Notice

Alexander Zoond or Alexander Sand and, sometimes, Alec Sand is one of my scientific heroes from the then new world of experimental biology. He is sometimes labelled as a comparative physiologists but what he did was excellent physiology; there is nothing to be gained by adding ‘comparative’ or demeaning its fundamental importance because it was not done on mammals. Only fairly recently have I got to know more about him and only last week discovered his tragic end.

Information on him comes mainly from his obituary notice for the Royal Society and a family tree (without much detail) on ancestry.com. F. S. Russell FRS (later Sir Frederick) wrote that Sand was ‘of Jewish-Russian extraction’ and was born in Warsaw on 28 December 1901. The unrest in Poland (his father was a Menshevik) saw the arrival of the family in London when Alec was six. Before Zoond the family name had been Zundelevich or, in South Africa, Zoondevitch.

His schooling in London was entirely on the arts side. After a year at Reading university college came a strange and unexplained move. Reading then was the place for degrees in agriculture but Zoond left for the University of British Columbia in Canada to study dairy bacteriology with the intention of a farming career. After graduating with an agriculture degree, he tried a job in farming in the U.S.A. for a year but found it unrewarding. In 1924 he headed back to Canada, this time to McGill University, for an MSc in bacteriology (his thesis from October 1925, The influence of green manures upon the growth and physiological efficiency of Azotobacter chroococcum, can be found in full online). Appointed to a Demonstratorship in 1926 he had a chance meeting with Lancelot Hogben FRS who was then Assistant Professor of Medical Zoology. Hogben was only at McGill for 18 months and when he was offered the chair of zoology at the University of Cape Town, he took it. With him went Zoond, having turned down a fellowship at Yale, as lecturer.

Lancelot Hogben
Photograph used in his Royal Society
Biographical Memoir

Zoond, in other words, changed field entirely, an attitude entirely in keeping with Hogben’s iconoclasm and loathing of classical zoology and classical zoologists. Hogben wrote:

…I must here explain that the sacred cow of first-year biology in those days was a whimsy of Thomas Henry Huxley known to posterity as the “type system”. This involved the detailed anatomical study of about a dozen species chosen from different groups of the Animal Kingdom, less because they were typical of their phyla or classes than because it was at one time relatively easy to place orders for them with pet shop dealers in London. When I entered my department on the day after arrival, I found the shelves lined with dozens of bottles of Huxleyan types imported in formalin from the seat of the empire. On enquiry, I learned that no living creatures had ever desecrated these hallowed precincts. The same day, I therefore instructed my laboratory steward to empty all the contents of the pickle jars in the yard behind the laboratory ready for the incinerator when the sun had dried them. Henceforth, the motto was to be only the living pass these portals…

Zoond clearly thrived in Cape Town. He published with Hogben on the pH of surface water, with David Slome¹ on the effect of electrolytes on cardiac rhythm and with Enid Charles, (the first wife of Hogben) on respiration in crabs.

In his memoir Hogben did not mention Zoond by name. However, it is clear who is being referred to in the following passage:

When I was not dealing face to face with the student body, Boer-backlashed mentality could make matters disagreeable. The lecturer whom I imported from McGill to replace the relict of my predecessor formed an attachment to one of the only two coloured medical graduates then practising in the Dominion, the other one being her brother-in-law, a leading light in the Moslem Community. Both held the Glasgow degree. When our young colleague confided in us that he was bringing his lady-friend to the University annual dance, both Enid and I scented embarrassment for both. Accordingly, we asked the pair to dine with us so that the lady could come as our own guest. My Jewish students clustered round and saw to it that she was never without a partner. Next day, the more rabid nationalist students held an open air campus meeting of protest.

It was after Hogben, with apartheid in the offing, had left Cape Town in 1930 that Zoond, who had stayed on, completed his work on chameleons, that on the tongue with Joyce Eyre², that on pigmentation with Eyre and with Naomi Adeline Helen Bokenham (later Millard) (1914-1997). The papers were published between 1933 and 1935, as he was leaving or after he had left South Africa.

Cape Dwarf Chameleon (Bradypodion pumilum, in the 1930s known as
Lophosaura pumila and then Microsaura pumila).
All of Zoond's (Sand's) research was on this species.
Photo: JonRicfield from here.

Sand (still Zoond then) left Cape Town in 1933; before doing so he took a Ph.D. from that university. He then went to Cambridge to work with James Gray FRS (later Sir James) then Reader in Experimental Zoology who was just beginning his famous work on animal locomotion. It was their joint work on the locomotory rhythm of the dogfish published in 1936 that was on ETBF’s reading list.

Alexander Zoond changed his name by deed poll on 9 March 1935 to Alexander Sand. The notice in the London Gazette gave his address as 9 Storey’s Way, Cambridge, his occupation as ‘biologist’ and stated that he was a naturalised British subject. I have not been able to work out in what capacity Sand was working with Gray. His second, this time Cambridge, Ph.D. was awarded in 1938. He must have been in statu pupillari to have been awarded a Ph.D. since those paid by the university were (and I think still are) disbarred. Did he have some form of external fellowship or did he fund himself?

In early 1935 he joined the staff of the Marine Biological Association’s famous laboratory at Plymouth as Physiologist. There he worked on the sensory physiology of fish including the lateral line, stretch receptors in the muscles and the ampullae of Lorenzini. The research on the auditory labyrinth he did with Otto Löwenstein (1906-1999, FRS 1955) was—and still is—of relevance in all vertebrates and clinical conditions. What happens in the semicircular canals during movement of the head was sorted out for the first time in experiments that would have been impossible in mammals.

Although I am not going to describe here the work on sense organs in fish, I thought it worth quoting the following paragraph that indicates the sensitivity of the lateral line and why Sand worked in the cellar of the Plymouth laboratory ‘which is hewn out of solid Devonian limestone’:

To demonstrate the delicacy of the response he formed a light hammer of a rubber stopper which was allowed to fall through a distance of 8 cm. on to a rubber sponge lying on the edge of the table about 3 feet away from the prepara­tion, the shocks imparted to the table being quite imperceptible to the human touch. 

There are a few traces in the newspapers of Sand’s time at Plymouth. The Western Morning News of 28 December 1936 reported that a fire had been discovered in Fardel Manor on the Cornwood-Ivybridge road which was occupied by Sand at the time. Large beams underneath a hearth in one of the upstairs rooms were smouldering. One of Sand’s children was due to sleep in the room. The fire brigade had to cut away the floorboards before extinguishing the fire ‘with the aid of patent extinguishers’. The medieval house, now Grade I listed and about 9 miles from the MBA laboratory, was once the seat of the Raleigh family.

By the time of the special census in 1939, the Sand family had moved a couple of miles to Gerston, Harford, now part of Ivybridge. Sand is shown there (as a research biologist) with his wife, Edith G Sand (born 19 April 1909), two children (records still closed) and Elizabeth B Reidy, born 1914, described as ‘paid companion help’. Russell wrote that he had two sons.

Motorists in Britain lived for decades in fear and dread of leaving their car without lights at night. Bored policemen pounding their beats were always on the lookout. Sand was fined ten shillings (50p) at Totnes Petty Sessions for ‘leaving a motor car without lights’ at Redworth Terrace, Totnes on 7 January 1940. How being fined for leaving a car without lights tied in with the wartime blackout regulations which demanded no lights, I do not know. Normal for Devon, maybe.

Russell then notes that Sand did all he could to get into the armed forces as quickly as possible after the outbreak of war.  He was commissioned on 1 May 1940 in Special Branch of the Royal Navy Volunteer Reserve. Promotion to Lieutenant followed in October 1940. He, like many biologists in the war, was involved in radar. Russell wrote:

After a time on the mainland in the north of Scotland he was posted as Officer in Charge of the station on the Shetlands. In 1941 Sand came south to lecture on radar for some months before being posted to a cruiser in which he was in the Indian Ocean. He contracted malaria in the Persian Gulf and when recovered went on leave to South Africa where his wife and two sons were living for the war years. He was then posted to a monitor and was present at the Sicilian and Salerno landings, his ship being hit. After a period of unemployment he was seconded for research at the Naval Department in the Medical Research Council’s Laboratory at Hampstead.

Russell went on to say that the research at Hampstead was on the physiology of diving and that in the course of his duties he received training in diving himself. It was, said, Russell, likely to remain confidential. G. L. Brown (FRS 1946, knighted 1957) was in charge of the research. The Physiological Society always maintained the tradition of using just initials and surnames in its internal communications, never honours or titles, so Sir George Lindor Brown FRS was always G.L. Brown. His biographical memoir contains more details of the research and includes Sand, along with a while string of well-known physiologists and naval officers, as members of the laboratory:

Brown discovered at the Lister Institute, and commandeered, the long-disused chamber that J. S. Haldane, the father of J. B. S., had used a generation earlier in the research that gave divers their first reliable decom­pression tables. The chamber was transported to the garden of the Institute at Hampstead, a hut was built around it, a new compressor was installed, and Brown and his associates began to acquire the somewhat esoteric expertise needed for doing human experiments under pressure. It was Brown’s first opportunity to recruit a research team of his own. H. P. Marks, Macintosh and Collison of the Hampstead staff were already available; they were joined by F. Dickens, C. B. B. Downman and 20-year-old H. B. Barlow; A. Sand and W. D. M. Paton arrived a little later. S. Gay French, H. M. Balfour and other naval medical officers, along with Ellis, were occasional members of the group; others who sometimes took part included Haldane, D. Williams and R. G. Bickford.

Sand was elected to the Royal Society on 16 March 1944.

But then came tragedy. on 11 July 1945 at 58 West End Lane, Hampstead, Alexander Sand shot himself in the heart with a pistol. The famous coroner, Sir Bentley Purchase recorded the verdict of the inquest on the death certificate, ‘dec[ease]’d did kill himself not being of sound mind’. I have been unable to find any press reports that might shed more light on the events leading to his death. There is no mention on the death certificate of service in the navy (occupation shown as .F.R.S. and Doctor of Science’). However he is listed as having died while still serving in the RNVR. His nominal ‘ship’ (all sailors are attached to a ship, even if that ship is a shore establishment) was H.M.S. Victory, the shore establishment at Portsmouth.

Russell concluded his obituary notice:

The whole Plymouth staff were anxiously looking forward to the day when they would once more be reunited at the laboratory and the blow was bitter when it was known that Alec Sand would not be numbered among those returning. Having reached the height of his research powers it was anticipated that he would open up new avenues in the little explored field of the sensory environment of marine animals. All alike felt keenly the loss of a friendly and inspiring colleague, and a gap has been created in the ranks of physiologists that it will be hard to fill. 

Thomas Alan Stephenson (1898-1961, FRS 1951), Lancelot Hogben’s successor in Cape Town until returning to Britain in 1940 as professor of zoology at Aberyswyth, introduced his obituary of Sand in Nature as follows:

In the completely unforeseen death of Alexander Sand at the age of forty-three, comparative physiology suffered a very severe loss. Not only was his later work distinguished in a remarkable degree, but so much more of the same calibre might have been expected from him in the next twenty years. 

UPDATED 11 October 2020

*Hogben claimed he had forgotten the name of his predecessor, ‘a dedicated necrophilist’. It was John Dow Fisher Gilchrist (1866-1926), a marine biologist born in Scotland, who, in his time, seems to have been far more modern in research, but not in teaching perhaps, than Hogben indicates.

1. David Slome (1906-1995). Details here.

2. I can find no further information on Joyce Eyre.

Alexander RMcN. 2001. Otto Egon Lowenstein. 24 October 1906 - 31 January 1999: Elected F.R.S. 1955. Biographical Memoirs of Fellows of the Royal Society 47, 357-368.

Hogben, LT. 1998. Lancelot Hogben. Scientific Humanist. Woodbridge, Suffolk: Merlin Press.

MacIntosh FC, Paton WDM. 1974. George Lindor Brown 1903-1971. Biographical Memoirs of Fellows of the Royal Society 20, 41-73.

Stephenson TA. 1945. Dr Alexander Sand. Nature 156, 383-384.

Russell FS. 1948. Alexander Sand 1901-1945. Obituary Notices of Fellows of the Royal Society 5, 803-815.

Wells GP. 1978. Lancelot Thomas Hogben 9 December 1895 - 22 August 1975. Elected F.R.S. 1936. Biographical Memoirs of Fellows of the Royal Society 24, 183-221.

Zoond, A. 1933. The mechanism of projection of the chameleon’s tongue. Journal of Experimental Biology 10, 174-185. 

Zoond A, Bokenham NAH. 1935. Studies in reptilian colour response. II. The role of retinal and dermal photoreceptors in the pigmentary activity of the chameleon. Journal of Experimental Biology 12, 39-43.

Zoond A, Eyre J. 1934. Studies in reptilian colour response. I. The bionomics and physiology of the pigmentary activity of the chameleon. Philosophical Transactions of the Royal Society B. 223, 27-55.