Sunday 30 July 2023

Red-bellied Conure or Maroon-bellied Parakeet: a colour plate from 1954

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 9th in the series.

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The committee that drew up the common names for birds did a terrible job on the parrots. Just about every small parrot with a long tale was called a parakeet from whatever part of the world it occurs. The word ‘conure’ for most of the South American species seems to have been dumped. Thus what was known as the Red-bellied Conure is now the Maroon-bellied Parakeet. Other common names, Brown-eared Conure, for example were commonly used by people who kept and bred the species—Pyrrhura frontalis. The species is found from southeastern Brazil to north-eastern Argentina, including eastern parts of Paraguay and Uruguay.

The article accompanying this plate was written by Arthur Alfred Prestwich (1903-1987) who was for many years Secretary of the Avicultural Society. He devoted a great deal of his time researching the history and first breeding of various species, but especially parrots, in captivity.

The plate was the work David Morrison Reid Henry (1919-1977) in 1955. He signed his work as D.M. Henry and was an artist favoured by the Avicultural Society for the plates published in this period.

Avicultural Magazine 60, 1954

Saturday 29 July 2023

The Curious Case of the Thirteen-lined Ground Squirrel: Digging in the History of Endocrinology

In a previous article I described my delight at seeing Thirteen-lined Ground Squirrels for the first time in the wild. These animals are a common sight in the grasslands and prairies of north America. My first encounter with this species was 58 years earlier. Before I joined him as a PhD student John Phillips (1933-1987) was back in Sheffield on his first long leave from the University of Hong Kong where he was Professor of Zoology, having been appointed three years earlier at the age of 29. In those days expat terms were such that after two years and seven months ‘home’ leave of five months was taken—an ideal arrangement for those wishing to do research. I found him in the attic, used as an animal house. He was concerned that a group of ground squirrels - kept together—was not thriving. I could see they were being fed on standard rat lab pellets plus the usual mixture of seeds, biscuit etc given to hamsters. I immediately suggested giving them more protein since I had kept a related species and knew that they ate insects and other invertebrates in addition to the typically rodent diet. The technician in charge  could not believe I was right but went off to collect a handful of mealworms which were thrown into the floor cage. The ground squirrels fell upon them and had soon scoffed the lot. With the problem of feeding solved, John finished his work on them and returned to Hong Kong. I assumed that the work had been published since in his report to the Nuffield Foundation in 1967, which funded his work in comparative endocrinology to the extent of pay, equipment and consumables for a research unit in Hong Kong, he listed among the publications one on the Thirteen-lined Ground Squirrel which was in preparation for submission to the Journal of Endocrinology. However, when I searched all his publications while researching this article I could not find it and only this week have I found that the work was never published and realised why it was never published

In the early decades of the 20th century, endocrinology was in its infancy and there was still argument, dating from the 1850s, of whether or not the adrenal glands are essential for life. The Thirteen-lined Ground Squirrel (now Ictidomys tridecemlineatus but then included in the genus Citellus and later Spermophilus) achieved its fame as an oddity in comparative endocrinology because it can consistently survive removal of the adrenal glands by virtue of the growth of adrenocortical tissue in the ovary. Richard Arnold Groat (1915-1994) was a PhD student in the Department of Anatomy at the University of Wisconsin when he published his findings in 1943 and 1944. Groat went on to work in a number of medical schools and in private practice as a pathologist in North Carolina.

Groat showed that in common with a number of other species, Thirteen-lined Ground Squirrels only survive after adrenalectomy if given 1% sodium chloride to drink. With that treatment the animals continue to feed. However, unlike in other species, Groat found that the salt water can be removed after a minimum of 15 days and that the ground squirrels can withstand a fast after 3-4 months. He then showed that adrenocortical-like tissue appeared in the ovaries after complete removal of the adrenal glands strongly indicating that adrenal steroid hormones were being produced by the ovary in quantities sufficient for normal life.

I do not know how Ian Chester Jones (1916-1996) heard of Groat’s work. There was no mention of it in his seminal monograph on the adrenal cortex which was published in 1957. However, he became sufficiently interested to start this work in 1961 at Harvard. He was again working as a visitor in Roy Orval Greep’s (1905-1997); he had spent a couple of years there in the 1950s on a Fellowship of the Commonwealth Fund of New York (later to become Harkness Fellowships). Ian Henderson, then a PhD student in Sheffield (1941-2019) was the second author of the paper published in 1963. They explained their interest in Groat’s work:

His [Groat’s] findings have always been open to the suspicion that they are merely manifestations of adrenal rests which showed particular responsiveness in this species. On the other hand, his work may indicate the capacity of ovarian tissue to change into a new type which may be adrenocortical both in form and function. Should this be so, then a question of fundamental importance is raised. It may be, for example, that the ground squirrel displays, in intensified fashion, a property that exists in mammals generally. For these reasons, this paper presents a re-examination of the problem.

In modern terms, there is another possible explanation: a population of pluripotent stem cells or partially differentiated cells in the ovary stimulated by ACTH from the pituitary to divide and differentiate into cells typical of the adrenal cortex rather than of the ovary, the ovary and adrenal cortex being of similar embryonic derivation. With the negative feedback between concentrations of adrenal steroid hormones and ACTH secretion, the concentrations of ACTH in the blood would have been sky high in the adrenalectomized animals.

Essentially Chester Jones & Henderson confirmed Groat’s findings. As with Groat, they were unable to find adrenal rests—small isolated clumps of adrenocortical tissue outwith the adrenal gland—in the normal adrenal. These occur in various locations including the ovary and testis in mammals.

The implication of the observations would be that with the apparent absence of adrenal rests, the ovary of the intact ground squirrel would not be producing adrenal steroid hormones. However, Gavin Vinson (1939-2021) then also working with Chester Jones in Sheffield found that ovarian tissue could convert progesterone to the adrenal steroid hormones, corticosterone. In his paper of 1965 he wrote (references omitted):

It is particularly interesting that the ovary has the capacity to convert progesterone to corticosterone. While this might be ascribed to adrenal rests, Groat (1943, 1944) and Chester Jones & Henderson (1963) were unable to find adrenocortical tissue in the ovaries and adnexia of normal ground squirrels. Another possibility is that ovarian cells, possibly the interstitium, of the ground squirrel are able to produce both sex and adrenocortical steroids. An attractive hypothesis is that, as the ovary shares a common embryological origin with the adrenal cortex, throughout the vertebrates both tissues are able (to a greater or lesser degree) to manufacture both oestrogens and corticosteroids.

While there is current molecular evidence against the latter in the latter explanation in mammals, I suspect the explanation possibly lies in a small population of partially differentiated adrenocortical cells in the ovary of the ground squirrel, i.e. cryptic and widely spread adrenal rests with the potential to grow into clumps of adrenocortical tissue.

William George Henry Seliger (1922-2016), A James Blair and Harland Winfield Mossman (1898-1991) followed up Groat’s work in 1966; they were in the same department as Groat in the University of Wisconsin and I suspect Groat may have been Mossman's student. They showed that the cells characteristic of the adrenal cortex which appear in the ovary are in close relationship to the epithelial remnants of the rete ovarii and the medullary cords. Identical cells were found around the efferent ductules of the testis in adrenalectomized males. Histochemical tests indicated the cells were identical to normal adrenocortical cells. In vitro, the areas of the ovary with the adrenal-like cells converted progesterone to adrenal steroid hormones identified as corticosterone and desoxycorticosterone (11-deoxycorticosterone)* the combination providing both glucocorticoid and mineralocorticoid activity. Confirmation of the vital role of the ovaries and testes in adrenalectomized ground squirrels was obtained:

Male and female animals adrenalectomized a year before and maintained on tap water died within four days when the gonads and adnexa were removed. Male animals adrenalectomized a year before and maintained on tap water survived actively for two weeks when the testicles were removed, provided that the efferent ductules and surrounding adrenal cortex-like cells were left in place. These animals died within four days after the adnexa were removed.

To return to the paper by Phillips and Chester Jones that was never published, the chronology suggests that the work they were doing in September/October 1965 produced results indistinguishable from those published by Seliger, Blair and Mossman in 1966. In short, they had been scooped.

Since the 1960s there has been little to add to the story. It would seem that cells with the potential  to become adrenal cortical cells can be retained during embryonic development in the ovaries and testes in the ground squirrel. In that respect there is nothing new—adrenal tissue has been found after adrenalectomy in similar locations of mesodermal origin in other mammals. In the rat a report from the 1920s indicated an incidence of 19%. Similarly, some human patients show accessory adrenal tissue after adrenalectomy. However, what makes the Thirteen-lined Ground Squirrel special is the extent and consistency of formation of adrenocortical tissue in the ovary and testis. The finding in 1967 of similar structures in the ovary after adrenalectomy of the Northern or Five-striped Palm Squirrel (Funambulus pennantii) by Seth and Prasad of the University of Delhi, suggests that sciurid rodents may all show the phenomenon.

The question remains of whether the ability to form new adrenocortical tissue has any functional significance. In other words, has the retention of mesodermal remnants capable of forming adrenocortical cells been subject to positive natural selection in some mammals—or are they simply vestiges that only assume a physiological significance under laboratory and clinical conditions that never occur in normal life?

A possible clue to some functional role of adrenocortical cells in the ovary comes from the rabbit. In 1974 Hiroshi Mori and Keishi Matsumoto reported:

Adrenocortex-like cells were consistently found in the mesovarium or in the hilus of the ovary of rabbits less than three months old. The adrenocortex-like cells occurred usually in the form of nodules among or near the mesonephric tubules. But they sometimes appeared within the rete ovarii either in direct contact with the epithelial cells or scattered in the stroma.

Histochemical studies indicated they were adrenocortical cells. Moreover, ovarian plus mesovarium tissue in vitro was able to convert progesterone to corticosterone. These studies, now 50 years old, raise another question: what is the role of adrenocortical hormones in the ovary of young rabbits?

If something similar obtains in the ovary of other mammals in early life, is is then possible that the squirrels have had functional adrenocortical cells in the ovary and/or testis earlier in life and that adrenalectomy simply stimulates the remaining population of dormant stem cells? Has anybody had a look?

*While the methods used to identify steroid hormones in the 1960s do not satisfy modern criteria, there is no reason to suspect they were not as stated by the various authors of the time.

Chester Jones I, Henderson IW. 1963. The ovary of the 13-lined Ground Squirrel (Citellus tridecemlineatus Mitchell) after adrenalectomy. Journal of Endocrinology 26, 265-262 doi:10.1677/joe.0.0260265

Groat RA. 1943). Adrenocortical-like tissue in the ovaries of adrenalectomized ground squirrel (Citellus tridecemlineatus). Endocrinology 32, 488-492 doi:10.1210/endo-32-6-488

Groat RA. 1944. Formation and growth of adrenocortical-like tissue in the ovaries of adrenalectomized ground squirrel. Anatomical Record 89, 33-41 doi:10.1002/ar.1090890104

MacFarland WE. 1945. The vital necessity of adrenal cortical tissue in a mammal and the effects of proliferation of cortical cells from dormant coelomic mesothelium. Anatomical Record 93, 233-249 doi:10.1002/ar.1090930303

Mori H, Matsumoto K. 1974. Constant occurrence of adrenocortical tissue in the juvenile rabbit ovary. American Journal of Anatomy 141, 73-89 doi:10.1002/aja.1001410105

Seliger W, Blair AJ, Mossman HW. 1966. Differentiation of adrenal cortex-like tissue at the hilum of the gonads in response to adrenalectomy. American Journal of Anatomy 118, 615-629 doi:10.1002/aja.1001180217

Seth P, Prasad MRN. 1967. Effect of bilateral adrenalectomy on the ovary of the five striped Indian palm squirrel, Funambulus pennanti (Wroughton). General and Comparative Endocrinology 8. 157-162. doi:10.1016/0016-6480(67)90124-4

Vinson GP. 1965. Steroid production in vitro by the ovaries and adrenal glands of the thirteen-lined ground squirrel (Citellus tridecemlineatus Mitchell). Jourbnal of Endocrinology 32, 401-402 doi:10.1677/joe.0.0320401

Wilson JB, Zopey M, Augustine J, Schaffer R, Chiang M, Friedman TC. 2021. High prevalence of adrenal remnant tissue in patients undergoing bilateral adrenalectomy for Cushing's disease. Hormones and Metabolic Research 53, 161-168 doi: 10.1055/a-1253-2854

Saturday 8 July 2023

Thirteen-lined Ground Squirrel

Never having seen the species in the wild I was delighted we had good views of a Thirteen-lined Ground Squirrel (Ictidomys tridecemlineatus) as it ventured out of its burrow to eat dandelion heads. We were in Des Lacs National Wildlife Refuge in North Dakota at the end of May.

Widely distributed in the grasslands and prairies, from Canada south to Texas, this ground squirrel was also known by a number of common names including Striped Gopher and Leopard Ground Squirrel. It is omnivorous taking insects and, apparently, even mice and shrews. I will return to its dietary requirements in another article since my knowledge of its omnivory apparently saved an experiment in the mid-1960s, earning a few soon-forgotten brownie points. These still images were taken from 4K video and processed in Photoshop and Topaz DeNoise AI.

Friday 7 July 2023

Flatters and Garnett Ltd. Microscopy and Biological Suppliers 1901-1967

I was surprised when I was looking up what had happened to Flatters & Garnett, the Manchester biological supplies firm that equipped school and elementary laboratories, as well as amateur microscopists, entomologists and botanists, in the first seven decades of the 20th century. I thought they must have been taken over as laboratory supply companies consolidated by mergers and takeovers. I was wrong. Flatters & Garnett went into liquidation in 1967 after what the Science Museum website described as ‘financial problems’.

Abraham Flatters (see my previous article here) was joined by Charles Garnett to form their partnership in 1901. The Garnett family after ‘a serious rift’ bought out Flatters in 1909 so Flatters & Garnett while retaining the name continued without Flatters.

Unfortunately, the Science Museum’s website does not state the sources of its information on Flatters & Garnett. Therefore I will try to flesh out some of the information provided.

Charles Garnett was born on 30 May 1842 at Latchford near Warrington, Cheshire, the son of a blacksmith. According to the Science Museum he became apprenticed to a grocer ‘but, at the age of 21, emigrated to New Zealand. There, he became interested in natural history and made a collection of the ferns found on South Island. In 1873, he returned to Manchester and opened a restaurant in Cateaton Street’. However, in the 1881 Census he is shown as a confectioner employing 3 girls and 1 boy. The same occupation is shown in 1891. Only in 1901 is he shown as ‘confectioner and restaurant keeper’. He had married in January 1876 in Manchester; his wife was born in Adelaide, South Australia. By 1911 he is just ‘confectioner’; there is no mention of his involvement with Flatters & Garnett. Also present were his wife, a son (a jeweller), his brother (retired blacksmith) and two servants. Charles Garnett died on 12 January 1921. He left £12,042 His executor was his son, John Benbow Garnett who was closely involved in Flatters & Garnett from the start.

John Benbow Garnett was born on 4 February 1877. He was educated at Ackworth, a Quaker school. In the 1901 Census he was living at home with his occupation shown as ‘pharmaceutical chemist’. He then joined Flatters in Flatters & Garnett and my guess is that his father provided the money while John worked with Flatters in the business. In the 1911 Census, John described himself as a ‘preparer of natural history objects and lantern slides’ work that was also being done by Flatters. Was it a clash between John and Abraham Flatters that led to the latter’s departure.

Advertisements from Flatters & Garnett in its first ten years show an emphasis first on the pharmacy and photographic activities. An advertisement ran in the Manchester City News (this one is from 1902) soon after Flatters & Garnett was founded:

Flatters & Garnett Ltd. Dispensing & Photographic Chemists, make a speciality of dispensing physicians’ prescriptions. “Neroline,” an emollient cream for chapped hands, 1s. and 1s.9Id. per bottle.-.48, Deansgate (one door from Blackfriars-st.).

By 1910, advertisements were appearing which emphasised a particular aspect of the business but by this time the pharmacy was not being mentioned.

Flatters & Garnett Ltd. Specialists in Photographic Work. 32 Dover Street (close to the University) Manchester S.E. Send for revised Price List, just issued. Developing Printing Enlarging. N.B.-No connection with any other firm.

This was in Manchester City News of 28 May 1910. The no connection with any other firm was of course to Abraham Flatters’s breakaway company that was subject to successful legal action by Flatters & Garnett in the same year.

The wording on 31 December 1910 was:

Lantern Slides. Large stock for sale or hire, or made to order from negatives, prints, drawings &c by experts.

For microscopy, this one appeared on 22 October 1910:

Microscopes, slides and accessories, for textile and other fibre examination, for serious work or recreation

By the time of the 1921 Census, John Benbow Garnett described himself as a pharmacist and Managing Director of Flatters & Garnett Ltd, scientific instrument makers. He led the expansion of the company and its establishment as a leading player in biological supplies.

This is from the Science Museum’s website:

Flatters and Garnett Ltd moved in 1913 to larger premises at 309 Oxford Road, opposite the University. About a year later, the company developed Mersol, an immersion oil for use with high power microscope objectives which became very popular and sold well for many years.

Flatters & Garnett Ltd expanded its business steadily during the 1920s. The company increased the range of instruments it produced, including dissecting microscopes and the Precision microprojector. The company won a reputation for producing well-designed, reliable instruments and sold its products all over the world. In 1932, the firm acquired a large Victorian house on Wynnstay Grove in Fallowfield where it moved the microslide, specimen, photographic and chemical departments. Here the staff could have more space and less disturbance from noise and dirt than on Oxford Road. In 1950, the company introduced the Mikrops industrial projector. This replaced the microscope for routine examination in many laboratories.

On 1 January 1938, Nature contained the following report:

A serious fire occurred on Sunday morning, December 19, at the laboratories of Messrs. Flatters and Garnett, Ltd., the well-known Manchester firm of microscopists. The chemical and microscopical laboratories were completely burnt out, but a considerable number of mounted slides were saved. The photographic and lantern slide department was only slightly involved and the new instrument workshop, for manufacturing microscopes, micro-projectors, etc., escaped entirely. All the staff is being retained, and work has already been resumed in temporary laboratories. Stocks of most of the firm's chemical specialities are held at the head office, 309 Oxford Road, Manchester, 13, which is two miles from the laboratories. 

Microscope Slide Label

The company continued to exhibit at the Manchester Microscopical Society. In 1949, for example, they demonstrated the ‘latest advances in microscopy’ at the annual public exhibition.

It seems likely that at some time John Benbow Garnett handed over the reins to his son, Wilfred John. On a sea trip to Australia in 1958-59 John Benbow is described as ‘microscopist’ which gives no real indication of whether he had retired or not.

John Benbow Garnett died on 10 January 1973, leaving £26,600. He had been living near Coleraine in Northern Ireland.

Wilfred John Garnett was born on 16 July 1914. In the 1939 Register, the emergency census, he was living at home and described as ‘Degree Biologist, Director of Laboratory’. It was Wilfred John who signed on 22 June the formal notice to wind up the company and to appoint liquidators:

…on Wednesday, the 14th day of June 1967, the following Extraordinary Resolution was duly passed: "That it has been proved to the satisfaction of this Meeting that the Company cannot by reason of its liabilities, continue its business, and that it is advisable to wind up the same and that the Company be wound up voluntarily…”

I wonder what had gone wrong. 

But that was not quite the end of the products. ‘A 1970 report of the Scottish Education Department on the school biology curriculum, stated: The transparencies of Marine Life taken by Dr. D.P. Wilson which were obtainable from Flatters and Garnett can now be purchased through W.J. Garnett, Breezemount, Ringrash, Macosquin, Coleraine, N.Ireland’. The address given was that occupied by Wilfred’s father at the time of his death.

Wilfred John Garnett died on 13 June 1988 in Bristol.

Flatters & Garnett Ltd produced a large number of catalogues throughout its existence. A number covered specific products, like stains or prepared microscope slides. The extent of their lantern slides stocks can be found in the catalogue which can be seen online here. The Second World War obviously left the company in some difficulty. Well into the 1950s the main catalogues had not been reprinted and an Interim Price List was published in 1951 and reprinted in 1954. Even then prices were not given; an accompanying leaflet did that.

As well selling as their own products, F&G were agents for Baker, Beck, Prior, Swift and Watson microscopes, for example.

The Science Museum in London contains a number of items made by Flatters & Garnett; see here.

Does any of the equipment and specimens supplied by Flatters & Garnett still survive in British schools? Microscope slides should have but I doubt they have. Do amateur entomologists still have F&G items still in use? Items do find their way onto eBay and into auction houses where the descriptions can provide great amusement. For example, a dissection kit is described as a ‘field surgical kit’ or a vasculum for botanists as a ‘metal satchel’. Amongst a number of cut-throat razors is one purportedly by Flatters & Garnett. I itch to tell the vendor that the F&G version was indeed a cut throat razor but only one side of the blade was hollow-ground (i.e. concave). Try shaving with a razor designed to cut sections of plant stems and the like by hand.

Flatters & Garnett Projection Microscope
From Micscape Magazine June 2002
see Microscopy-UK website
I have seen one of these lying around but
I cannot remember where

A Flatters & Garnett microscope for
sale on eBay at present

1954 reprint of 1951 catalogue

Microscope slide cabinet

Vasculum for plant collectors - to keep specimens
fresh. Completely replaced by the polythene bag