Salt Glands Revisited. The tragic story of an early salt-gland researcher and his family

Fifty years ago the late Jim Linzell and I were writing our monograph, Salt Glands in Birds and Reptiles, for the Physiological Society's Monograph Series; it was published in May 1975. In this series I revisit some of the topics and people who followed up the discovery of salt glands in birds by Knut Schmidt-Nielsen.

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Larry Zane McFarland
from Find a Grave

As well as Hubert and Mable Frings and the albatrosses of Midway (here) others followed up the discovery of salt glands by seeing which other species had them and how they worked. One was Larry Zane McFarland of the Department of Veterinary Anatomy at the University of California at Davis. He had a series of short notes in Nature in the late 1950s and 1960s (when that journal published letters of wide-ranging and topical interest). Amongst them was one which confirmed and extended the finding in albatrosses that ‘excitement’ induced or enhanced secretion. He worked on gulls and when producing secretion in response to a salt load found that light or sound increased the rate. The rapidity of response to shining a light at the birds or playing ‘individual notes, chords or a melody’ suggests the involvement of higher centres in the brain modulating the secretory reflex. McFarland described the musical instrument he played to the birds as a ‘quittar’, an old English word still used apparently and particularly in the USA for ‘guitar’.

In 1964 McFarland also introduced use of the ‘minimal stimulatory load’, the quantity of salt that needs to be introduced into the blood circulation in order to turn the salt glands on. That enable me to investigate the short-term and medium-term stimulatory and inhibitory effects of hormones and drugs on salt-gland secretion. 

Only after publication of our Salt Glands in Birds and Reptiles did I hear what had happened. I was at a conference in the USA, talking to someone who had worked at the vet school at Davis. I casually mentioned that McFarland, whom I had been quoting in the book, worked there. I was taken aback by the response that he was a dreadful man who had killed his wife and family, and then himself. And that is all I knew until a couple of weeks ago.

from Peaker & Linzell 1975

Larry Zane McFarland was born on 16 December 1930 in Des Moines, Iowa. There he attended Roosevelt High School. Before the veterinary course at Davis he was a student at Berkeley. He was known as a very hard worker. To fund his studies he worked a 40-hour week on night shifts.

He was awarded the DVM degree in 1956 and a PhD in 1960. At Davis he worked his way up to become chairman of the veterinary anatomy department, where he was remembered as an excellent teacher, in addition to doing research on a variety of subjects.

Hard working as he was, a colleague noted: ‘He used to preach to us that he did not expect us to devote too much extra time at the university, that it should go to our families. His devotion to his family was complete and total’.

McFarland had bought a large house with land in Davis. And it was there on 8 April 1972, that firemen found the bodies of McFarland, his wife Sonia, and their three children, Michael (14), Kenneth (10) and Nina (9), in the burning building. The two boys and McFarland had been shot while Sonia and Nina had been bludgeoned to death. A note was found in his vehicle containing the statement, ‘I am taking the life of myself and my family in the belief that a family should remain together in life or in death”.

The proximate cause of his behaviour that night seems to have been a phone call from his wife earlier in the day (7 April). Unbeknownst to most of his colleagues Larry and Sonia had become estranged, with Larry having lived in a nearby apartment for several months. 

Although a colleague in the department described McFarland as outgoing in personality he said that McFarland, like his colleagues, ‘led a "lonely existence" because of a "peer contact-peer judgment system of promotion" on university campuses. This "lonely existence" prevented any of his associates from detecting that something was going wrong. He said the problems McFarland was experiencing may have been deliberately kept secret as "a factor of his life which might have affected our opinion of him and influenced our decisions regarding his possible advancement”. The work situation common to many campuses, he said, "prohibits human exchange of human concern of one man for another”’.

The physical ramifications of McFarland’s actions that night, were evident in Davis for decades. The house McFarland had bought was the 15-room Chiles Mansion, built in 1902, which also had a large barn and other outbuildings on five acres. The house was gutted by the fire which McFarland had started with petrol in four or five places.

After a legal battle, Sonia’s mother gained the right to go against Larry’s wishes for the family to be buried together and she also gained possession of the property. The remains of the house were left untouched until McFarland’s mother-in-law died in 2006, aged 98. The barn was used a livery stable and tales can be found on the internet of the place being haunted, with horse owners reporting mysterious movements of the animals and the sighting of ephemeral white figures. The estate was sold to developers for $4.2M in the late 2000s and is now the site of 90-odd new houses on ‘Chiles Ranch’.

Thus behind the bland list of references, McFarland (1964) etc, at the end our of book, and in other papers, there is nothing that prepares anybody for the utter horror of McFarland’s actions in April 1972. All we can do is remember that Michael would now have been 66, Kenneth 62, and Nina 61.

The Argus, Fremont, California

Grey-headed Kingfisher: a colour plate from 1960

In the days when colour printing was extremely expensive, the Avicultural Society had special appeals for funds to support the appearance in Avicultural Magazine of the occasional colour plate. A well-known bird artist was then commissioned. Although the whole run of the Society’s magazines can be found online, the plates rarely see the light of day. Therefore I decided to show one, now and again, on this site. This is the 20th in the series.

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The artist for this plate was Chloe Elizabeth Talbot Kelly (born 1927) who went on to illustrate a number of field guides. Her paintings of birds appear in art sales. She began painting in 1945 at the Natural History Museum in London.

The article accompanying the plate was written by Alan Reece Longhurst (1925-2023). He was a well-known oceanographer and expert on plankton communities who spent a short time working in fisheries in New Zealand. He was born in Plymouth and after four years in the army he returned to London and university life. He graduated in entomology and then proceeded to a PhD on the ecology of notostracans. Fisheries research in West Africa then followed (with the short period in New Zealand in the middle). Spells in Plymouth and the USA were followed by a career in Canada. He became Director-General of the Bedford Institute of Oceanography in Nova Scotia.

The Grey-headed Kingfisher (Halcyon leucocephala) is widely distributed in Africa. Like many kingfishers it does not live up to its name. It is terrestrial, never diving for aquatic prey, feeding on Insects, other invertebrates, mice; lizards, frogs etc.. It does dive into water but only to bathe. 

Avicultural Magazine 66, 1960

Salt Glands Revisited. Hubert and Mable Frings in 1958: The Salt Glands of the Albatross

Fifty years ago the late Jim Linzell and I were writing our monograph, Salt Glands in Birds and Reptiles, for the Physiological Society; it was published in May 1975. In this series I revisit some of the topics and people who followed up the discovery of salt glands in birds by Knut Schmidt-Nielsen.

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Knowledge of how birds survive at sea can be divided into two eras: Pre-1957 and Post-1957. It was in 1957 that Knut Schmidt-Nielsen (1915-2007), along with his colleagues, Carl Christian Barker Jörgensen and Humio Osaki, who were visiting scientists at Duke University in North Carolina, discovered salt glands. These organs secrete salt at very high concentrations allowing marine birds to get rid of excess salt which they may ingest by drinking seawater and/or by eating invertebrates like squid which contain much more salt than fish.

The kidneys of birds and reptiles, in contrast to those of mammals, simply cannot produce urine with a high salt concentration. Therefore the discovery of an extrarenal route of salt excretion from glands in the head and emptying into the nostrils explained why marine birds could drink water and survive on salt-rich prey containing. Until then there had been no agreement, some observers claiming that not only could such birds drink seawater but also that, in captivity, seawater was essential for their survival.

Few zoos attempted to keep pelagic birds—those that remain far out at sea for months or years. But those that did came to the conclusion that they must be provided with saltwater or with salt added to their food, usually fish. In 1925 Glover Morrill Allen (1879-1942) then recently appointed to lecture in zoology at Harvard, wrote in his book, Birds and Their Attributes of 1925:

Most sea-birds are known to drink salt water in preference to fresh; indeed captive gulls may die without it.

That is why, my previous post on the keeping and failing to keep Waved Albatrosses at Brookfield Zoo, Chicago, in 1935, is interesting. They were fed by hand on freshwater fish dipped into salty water.

However, the work of Schmidt-Nielsen and his colleagues threw no light on why pelagic birds would need salt water in order to live. In all species of birds, including domestic ducks and geese, tested thus far, the salt glands only turn on in response to the ingestion of salt water sufficient to raise the osmolarity of the blood. No salt water: no secretion. Thus pelagic birds should be fine living on fresh water. It was only several months after the full paper by Schmidt-Nielsen and his colleagues was published that this apparent anomaly came to light.

Schmidt-Nielsen’s discovery had actually started in the late 1930s but the war and his move to the USA had delayed his following up his earlier, inconclusive, experiments in Norway. The discovery was followed up by reports from those who had realised they had seen salt gland secretion in actioon, drops of fluid appearing on the end of beaks for example, without appreciating the significance. Others with access to marine birds, followed suit to demonstrate the presence of salt glands in other species. 

In January 1958, Hubert and Mable Frings of the Department of Zoology and Entomology at Penn State University were on Midway, that famous island in the Pacific. They worked on sound communication in insects (Hubert had started off as, primarily, an entomologist, and were trying to see if sound could be used in pest control, invertebrate and vertebrate. It would appear they were on Midway because of a contract they had or were about to get with the US Office of Naval Research. The US Navy was very concerned because aircraft and albatrosses have a habit of colliding. In a report of research done between 1959 and 1963, the scale of the problem can be seen:

At Midway Naval Station, 1,100 miles west-northwest of Honolulu, military aircraft collide with flying albatrosses at the rate of about 300 to 400 per year. One aircraft out of every five that hits an albatross on takeoff either aborts (stops before it is airborne), or dumps fuel and returns for appraisal of damage. About 70,000 pairs of Laysan albatrosses [Phoebastria immutabilis] and 7,000 pairs of blackfooted albatrosses [Phoebastria nigripes] nest at Midway in any given year. The population is declining.

The Frings knew of the then very recent discovery of salt glands and took the opportunity to look for any signs of salt glands in action:

The birds were, therefore, observed with this in mind, and it was noted that after fighting among the Black-foots, or occasionally during the ritual dancing, some of the birds showed drops of fluid at the tips of the beaks. Furthermore, during a local harassment campaign, many of the birds in the treated area were actively dripping. This suggested that the gland could be activated by “stress” or excitement.

There was a limit to what could be achieved on Midway and so the US Navy agreed to fly four birds of each of the two species to Pennsylvania. They arrived on 18 April and were housed initially in an old hen house on the university farm. The vicissitudes of getting the birds to swallow the offered food and unsatisfactory housing and water supplies were explained in a full paper in Condor, and here I will only describe some salient points. The Black-footed Albatrosses soon learnt to take food offered by hand. The Laysan did not and until it was found that a piece of fish, for example, could be slid into the beak nearer the mouth and would be swallowed, they declined. Two of the latter died but not of inanition but, it would seem, a lack of salt. Handling and disturbance was seen to result in secretion by the salt glands but the birds drank the fresh water provided. When the two remaining Laysans were declining badly, it was decided to give each an injection of salt. Recovery was remarkably rapid. Within ten minutes the birds were up and walking normally. The Fringses concluded that because salt-gland secretion can be induced by excitement or “stress” in these birds and because they had no opportunity to drink salt water, they became hyponatraemic. They seemed unable to distinguish fresh water from salt water.

These problems were overcome by providing salt in gelatine capsules embedded in the fish used to feed the birds. Later, an artificial  sea water pool was provided and the birds were seen drinking from it.

The remaining birds thrived and were moved by air, first in June  to the Mount Desert Island Biological Laboratory, Salisbury Cove, Maine, where they were given real sea water in tubs from which they drank. Then, on  20 August 1958, they were sent to the  National Zoological Park in Washington, D.C. ‘where Drs. Theodore Reed and Malcolm Davis are continuing with the methods described here’. I do not know what became of the birds after they reached Washington.

Thus Hubert & Mable Frings explained why some marine birds need salt water to drink when in captivity. In the wild any loss of salt evinced by the salt glands switching on seemingly inappropriately would easily be made up from squid in the diet or by drinking sea water.

I will return in a later article to the question of salt-gland secretion appearing or the rate being changed by internal or external stimuli caused other than by an increase in the tonicity of the blood plasma. I will do so with some trepidation since the story ends with a most horrific human tragedy.

The main reason for the birds being caught and flown to Pennsylvania was of course to investigate the salt-gland secretion. This was done while the albatrosses were still at Penn State. For this phase, Hubert Frings enlisted the collaboration of Adam Anthony from the physiology section of the Biology Department (1923-2012) and Martin Warren Schein (1925-1998) who had wide interests in animal behaviour and then in the Poultry Science Department. Their paper was published in Science. When stimulated by supplying salt capsules in the food, the secretion contained between 792 and 856 m-moles of sodium per litre. This concentration of sodium is twice that of sea water.

The concentration of salt-gland secretion compared with sea water is of course the key to understanding the role of the salt gland in marine birds. While it is convenient shorthand to describe the function of the glands as getting rid of excess salt, it would be of no use to the bird if the concentration did not exceed that of sea water. The whole mechanism is there because it enables birds to obtain osmotically-free water. As such it is a highly effective desalination plant that requires some of the highest rates of blood flow ever observed while its cells are stuffed with mitochondria to drive the energetically expensive transport systems across its cell membranes.

Although the rate of secretion was not measured the group assembled by Hubert Frings noted the secretion is discharged from the nasal glands through small openings below each tube-nostril. The fluid then flows along grooves on the beak to drip off the end. After being given a salt load, drops were seen falling at internals of 2-4 seconds.

A Northern Royal Albatross (Diomedea sanfordi) at the Royal Albatross Centre, Dunedin, New Zealand
I photographed this one through the glass of the hide in 2019. The small openings below the tube-nostrils
together with the grooves along the beak can be seen clearly.

But who were the Fringses who were in the right place and at the right time to embark on a study of the albatross salt gland?

Hubert William Frings was born in Philadelphia on 1 January 1914. His father, born in Germany, was a flooring contractor. Hubert graduated from what became Pennsylvania State University in 1936. He received a Master’s degree from the  University of Oklahoma in 1937, and a PhD from the University of Minnesota, in 1940. Teaching jobs, traceable to Iowa and Missouri, can be found in the records. He became an insect physiologist at the US Army’s Chemical Center in Maryland. He taught biology for a short time at Gustavus Adolphus College, Saint Peter, Minnesota in 1946-1947 but then returned to Penn State where he climbed the academic ladder until 1961. Then he went to the University of Hawaii until 1966. Hubert and Mable are commemorated there by an endowed scholarship. Finally, he joined the University of Oklahoma, becoming David Ross Boyd professor until his retirement in 1979.

Mable and Hubert Frings on their wedding
day, 9 June 1936
(from a family tree on ancestry.com)

Hubert married Mable Ruth Smith on 9 June 1936. Her family were farmers in Pennsylvania. She was born on 11 April 1912.

Hubert and Mable wrote two books: together: Animal Communication in 1964 and Concepts of Zoology in 1970.

As part of their support from the US Office of Naval Research, Hubert and Mable, together with their son Carl Frederick, published An Annotated Bibliography on North Pacific Albatrosses in 1966.

Mable died on 28 March 1998; Hubert on 8 September 2008, both in New Hampshire.

…Meanwhile on Midway the albatrosses are thriving with over 2 million birds nesting on 2.4 square miles of the atoll. The US Navy left in 1993 and flights to and from Henderson Field only occur at night during the nesting season.

Allen GM. 1925.  Birds and Their Attributes. Boston: Marshall Jones.

Frings H, Anthony A, Schein MW. 1958. Salt excretion by the nasal gland of Laysan and black-footed albatrosses. Science 128, 1572.

Frings H, Frings M. 1959. Observations on salt balance and behavior of Laysan and black-footed albatrosses in captivity. Condor 61, 305-14.

Peaker M, Linzell JL. 1975. Salt Glands in Birds and Reptiles. Cambridge: Cambridge University Press.

Waved (Galapagos) Albatross at Brookfield Zoo, Chicago, in 1935: a colour plate from 1969

In the days when colour printing was extremely expensive, the Avicultural Society had special appeals for funds to support the appearance in Avicultural Magazine of the occasional colour plate. A well-known bird artist was then commissioned. Although the whole run of the Society’s magazines can be found online, the plates rarely see the light of day. Therefore I decided to show one, now and again, on this site. This is the 19th in the series.

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The artist of this plate and author of the accompanying article was Karl Plath (1886-1970). Plath had trained and exhibited as an artist in Chicago before being asked to become Curator of Birds at what is now called Brookfield Zoo in 1935; he retired in 1961. He also had a large private collection of birds.

Plath describes how three Waved or Galapagos Albatrosses (Phoebastria irrorata) were kept at Brookfield Zoo ‘some years ago’. One died after seven months, another after ten. But one lived for 4 years and six months—not long in albatross terms but considered a success at the time. Unable to take off in the large flight cage (30 x 12 metres and 7 metres high) the bird, thought to be a male, walked around swinging its head from side to side.

Plath does not provide a date but I have found a press photograph for sale online which shows two of the birds and is dated 25 July 1935, i.e. shortly after Plath joined the zoo. Who collected the birds—and why?

This date of 1935 means there is more to this article than a simple description of events because the way in which the bird was fed at the time is of interest to those of us who have been involved in the physiology of marine birds and the history of a key discovery in this field in the 1950s. I will expand on this thread in a future article.

Avicultural Magazine 75, 1969

The Besra in Hong Kong

This Besra (Accipiter virgatus) was photographed a few weeks ago in a tree it sometimes visits in the morning. The Besra is a forest bird but is known to frequent perches in the open at that time of day. It is widely distributed in southern Asia.

It is only in recent decades that the taxonomy and identification of sparrowhawks has been worked out. In the 1960s the Besra was completely unknown to Hong Kong birders. That is because there were only though to be two species visiting Hong Kong: the Japanese Sparrowhawk (Accipiter gularis) and the Eurasian Sparrowhawk (Accipiter nisus). Both were thought to be passage migrants.  The Japanese Sparrowhawk and the Besra were thought to be geographical variants of the same species. When it was realised that two different species were involved, and it was found that one is actually resident in Hong Kong (Besra) and the other entirely migratory (Japanese) did the position become clear. The Eurasian is still a rare migrant.

Identification of the Besra as a Hong Kong bird, also cleared up our scratching of heads in earlier years, starting in 1966,  as we saw display flights of sparrowhawks above Hong Kong. We could not decide between the two stated to be there at the time. They seemed too big for Japanese and not in the season for migrants of either species. And would migrants be displaying?

The Besra must have become more common as there are more and more trees and bushes compared with the hillsides devastated for decades by the collection of firewood for cooking during the Japanese occupation. The latest field guide has them as ‘fairly common, widespread resident and possibly autumn migrant’ whereas in 2005 it was down as ‘a scarce resident, possibly with increased numbers in autumn’.

What’s this moth we saw in Portugal?

 

In a wood near Aljezur we saw this moth, best described as very hairy with very impressive antennae. Only after we got back home did we non-lepidopterists track down what it was. To confuse matters, it has recently had its name changed. We found a photograph online, also from Aljezur, of Ptilocephala albida. However, that species has been ‘split’ recently and the form in Portugal is now Ptilocephala lorquiniella.

The species is psychid moth, known collectively as bagworm moths since the larvae construct from silk and debris a case in which the rest and then pupate. Only adult males fly.

The Medawar 1959 Reith Lectures, ‘The Future of Man’

Soon after I had started ‘A’ levels at school in 1959, the two of us doing zoology were joined at the door of biology block by Jim Key (James John Key 1917-1976) the senior biology master. He had been arguing with Stan Revill (1907-1993), senior history master, and was keen to test his line for further sparring in the staff room. They had both listened to the BBC’s Reith Lectures by P.B. Medawar entitled The Future of Man. That was, of course, before Medawar had become Sir Peter and had been awarded the Nobel Prize. The staff room argument was about whether natural selection in Man, was over (Stan Revill’s stance) or not (Jim Key’s).

Fast forward from 1959 to 2024, I realised that I had never heard these lectures and thus found the one available on the BBC website (here). I also found a copy of the book Medawar published immediately after the lecture series. I think, having read virtually all of Medawar’s later books and having written on his insights in endocrinology, my expectations were too high since I was disappointed both by the lecture (the sixth of the six broadcast on 20 December 1959) and by the book. Was I judging with the benefit of knowledge gained over the past 65 years? Or were his choice of topics,  analogies and explanations not really that good? I strongly suspect that even for his audience on the Third Programme on a Sunday evening, his presentations would have gone over the heads of most.

At the same time as reading Medawar of 1959, I have been reading Venki Ramakrishnan’s Why We Die of 2024. I found myself agreeing with his views on Medawar’s later works:

Unlike many scientists who focus narrowly on one area, Medawar, like Haldane, had widespread interests, and wrote books that were famous for their erudition and elegant writing. Many scientists of my generation grew up reading his Advice to a Young Scientist (1981), which I found pompous, arrogant, thoughtful, engaging, and witty all at once.

The Future of Man can perhaps be described as a series of early essays, in the way of early works by an artist. The best was yet to come. There were though patches of where he put his message over extremely well, as on the genetic consequences of ‘social habit or act of legislation’ and of assortative mating. The book of the lectures is historically important because it deals with topics of human biological and cultural evolution that were being talked about by those in Medawar’s circle in the 1950s. I was therefore surprised to see that my cheap copy was one which had been withdrawn from a British university library. Do such institutions now just throw away important books by important authors away? I fear we are already depriving students the pleasures of browsing in a proper library and then wondering why they are bright but not learned.

…and can you imagine a publisher or the BBC producing anything with that title, or including the word ‘eugenics’ in the text, in 2024?

Medawar PB. 1959. The Future of Man. London: Methuen.