The Salt Glands of the Albatross


Fifty years in the past the late Jim Linzell and I have been writing our monograph, Salt Glands in Birds and Reptiles, for the Physiological Society; it was printed in Might 1975. On this collection I revisit a number of the matters and individuals who adopted up the invention of salt glands in birds by Knut Schmidt-Nielsen.

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Data of how birds survive at sea may be divided into two eras: Pre-1957 and Submit-1957. It was in 1957 that Knut Schmidt-Nielsen (1915-2007), alongside along with his colleagues, Carl Christian Barker Jörgensen and Humio Osaki, who have been visiting scientists at Duke College in North Carolina, found salt glands. These organs secrete salt at very excessive concentrations permitting marine birds to do away with extra salt which they could ingest by ingesting seawater and/or by consuming invertebrates like squid which comprise rather more salt than fish.

The kidneys of birds and reptiles, in distinction to these of mammals, merely can not produce urine with a excessive salt focus. Subsequently the invention of an extrarenal route of salt excretion from glands within the head and emptying into the nostrils defined why marine birds may drink water and survive on salt-rich prey containing. Till then there had been no settlement, some observers claiming that not solely may such birds drink seawater but in addition that, in captivity, seawater was important for his or her survival.

Few zoos tried to maintain pelagic birds—those who stay far out at sea for months or years. However those who did got here to the conclusion that they have to be supplied with saltwater or with salt added to their meals, normally fish. In 1925 Glover Morrill Allen (1879-1942) then not too long ago appointed to lecture in zoology at Harvard, wrote in his ebook, Birds and Their Attributes of 1925:

Most sea-birds are recognized to drink salt water in place of recent; certainly captive gulls could die with out it.

That’s the reason, my earlier submit on the preserving and failing to maintain Waved Albatrosses at Brookfield Zoo, Chicago, in 1935, is attention-grabbing. They have been fed by hand on freshwater fish dipped into salty water.

Nevertheless, the work of Schmidt-Nielsen and his colleagues threw no mild on why pelagic birds would wish salt water with the intention to reside. In all species of birds, together with home geese and geese, examined to this point, the salt glands solely activate in response to the ingestion of salt water ample to boost the osmolarity of the blood. No salt water: no secretion. Thus pelagic birds ought to be wonderful dwelling on recent water. It was solely a number of months after the total paper by Schmidt-Nielsen and his colleagues was printed that this obvious anomaly got here to mild.

Schmidt-Nielsen’s discovery had truly began within the late Nineteen Thirties however the conflict and his transfer to the USA had delayed his following up his earlier, inconclusive, experiments in Norway. The invention was adopted up by reviews from those that had realised they’d seen salt gland secretion in actioon, drops of fluid showing on the top of beaks for instance, with out appreciating the importance. Others with entry to marine birds, adopted go well with to exhibit the presence of salt glands in different species. 

In January 1958, Hubert and Mable Frings of the Division of Zoology and Entomology at Penn State College have been on Halfway, that well-known island within the Pacific. They labored on sound communication in bugs (Hubert had began off as, primarily, an entomologist, and have been making an attempt to see if sound may very well be utilized in pest management, invertebrate and vertebrate. It could seem they have been on Halfway due to a contract they’d or have been about to get with the US Workplace of Naval Analysis. The US Navy was very involved as a result of plane and albatrosses have a behavior of colliding. In a report of analysis accomplished between 1959 and 1963, the size of the issue may be seen:

At Halfway Naval Station, 1,100 miles west-northwest of Honolulu, navy plane collide with flying albatrosses on the price of about 300 to 400 per yr. One plane out of each 5 that hits an albatross on takeoff both aborts (stops earlier than it’s airborne), or dumps gasoline and returns for appraisal of harm. About 70,000 pairs of Laysan albatrosses [Phoebastria immutabilis] and seven,000 pairs of blackfooted albatrosses [Phoebastria nigripes] nest at Halfway in any given yr. The inhabitants is declining.

The Frings knew of the then very current discovery of salt glands and took the chance to search for any indicators of salt glands in motion:

The birds have been, due to this fact, noticed with this in thoughts, and it was famous that after combating among the many Black-foots, or sometimes in the course of the ritual dancing, a number of the birds confirmed drops of fluid on the ideas of the beaks. Moreover, throughout an area harassment marketing campaign, lots of the birds within the handled space have been actively dripping. This steered that the gland may very well be activated by “stress” or pleasure.

There was a restrict to what may very well be achieved on Halfway and so the US Navy agreed to fly 4 birds of every of the 2 species to Pennsylvania. They arrived on 18 April and have been housed initially in an previous hen home on the college farm. The vicissitudes of getting the birds to swallow the provided meals and unsatisfactory housing and water provides have been defined in a full paper in Condor, and right here I’ll solely describe some salient factors. The Black-footed Albatrosses quickly learnt to take meals provided by hand. The Laysan didn’t and till it was discovered {that a} piece of fish, for instance, may very well be slid into the beak nearer the mouth and can be swallowed, they declined. Two of the latter died however not of inanition however, it could appear, a scarcity of salt. Dealing with and disturbance was seen to end in secretion by the salt glands however the birds drank the recent water offered. When the 2 remaining Laysans have been declining badly, it was determined to present every an injection of salt. Restoration was remarkably speedy. Inside ten minutes the birds have been up and strolling usually. The Fringses concluded that as a result of salt-gland secretion may be induced by pleasure or “stress” in these birds and since they’d no alternative to drink salt water, they turned hyponatraemic. They appeared unable to differentiate recent water from salt water.

These issues have been overcome by offering salt in gelatine capsules embedded within the fish used to feed the birds. Later, a man-made  sea water pool was offered and the birds have been seen ingesting from it.

The remaining birds thrived and have been moved by air, first in June  to the Mount Desert Island Organic Laboratory, Salisbury Cove, Maine, the place they got actual sea water in tubs from which they drank. Then, on  20 August 1958, they have been despatched to the  Nationwide Zoological Park in Washington, D.C. ‘the place Drs. Theodore Reed and Malcolm Davis are persevering with with the strategies described right here’. I have no idea what turned of the birds after they reached Washington.

Thus Hubert & Mable Frings defined why some marine birds want salt water to drink when in captivity. Within the wild any lack of salt evinced by the salt glands switching on seemingly inappropriately would simply be made up from squid within the eating regimen or by ingesting sea water.

I’ll return in a later article to the query of salt-gland secretion showing or the speed being modified by inside or exterior stimuli prompted aside from by a rise within the tonicity of the blood plasma. I’ll achieve this with some trepidation for the reason that story ends with a most horrific human tragedy.

The principle purpose for the birds being caught and flown to Pennsylvania was after all to research the salt-gland secretion. This was accomplished whereas the albatrosses have been nonetheless at Penn State. For this part, Hubert Frings enlisted the collaboration of Adam Anthony from the physiology part of the Biology Division (1923-2012) and Martin Warren Schein (1925-1998) who had extensive pursuits in animal behaviour after which within the Poultry Science Division. Their paper was printed in Science. When stimulated by supplying salt capsules within the meals, the secretion contained between 792 and 856 m-moles of sodium per litre. This focus of sodium is twice that of sea water.

The focus of salt-gland secretion in contrast with sea water is after all the important thing to understanding the function of the salt gland in marine birds. Whereas it’s handy shorthand to explain the operate of the glands as eliminating extra salt, it could be of no use to the chook if the focus didn’t exceed that of sea water. The entire mechanism is there as a result of it allows birds to acquire osmotically-free water. As such it’s a extremely efficient desalination plant that requires a number of the highest charges of blood circulation ever noticed whereas its cells are filled with mitochondria to drive the energetically costly transport programs throughout its cell membranes.

Though the speed of secretion was not measured the group assembled by Hubert Frings famous the secretion is discharged from the nasal glands by small openings under every tube-nostril. The fluid then flows alongside grooves on the beak to drip off the top. After being given a salt load, drops have been seen falling at internals of 2-4 seconds.

A Northern Royal Albatross (Diomedea sanfordi) on the Royal Albatross Centre, Dunedin, New Zealand
I photographed this one by the glass of the disguise in 2019. The small openings under the tube-nostrils
along with the grooves alongside the beak may be seen clearly.

However who have been the Fringses who have been in the best place and on the proper time to embark on a examine 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 turned Pennsylvania State College in 1936. He acquired a Grasp’s diploma from the  College of Oklahoma in 1937, and a PhD from the College of Minnesota, in 1940. Educating jobs, traceable to Iowa and Missouri, may be discovered within the data. He turned an insect physiologist on the US Military’s Chemical Heart in Maryland. He taught biology for a short while at Gustavus Adolphus Faculty, Saint Peter, Minnesota in 1946-1947 however then returned to Penn State the place he climbed the educational ladder till 1961. Then he went to the College of Hawaii till 1966. Hubert and Mable are commemorated there by an endowed scholarship. Lastly, he joined the College of Oklahoma, changing into David Ross Boyd professor till his retirement in 1979.

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

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

Hubert and Mable wrote two books: collectively: Animal Communication in 1964 and Ideas of Zoology in 1970.

As a part of their assist from the US Workplace of Naval Analysis, Hubert and Mable, along with their son Carl Frederick, printed An Annotated Bibliography on North Pacific Albatrosses in 1966.

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

…In the meantime on Halfway the albatrosses are thriving with over 2 million birds nesting on 2.4 sq. miles of the atoll. The US Navy left in 1993 and flights to and from Henderson Area solely happen at night time in the course of 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 steadiness and habits 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 College Press.

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