Vets and Huskies in the Antarctic – Tony Palmer (continued)

VETS AND HUSKIES IN THE ANTARCTIC – Tony Palmer (continued)

The purpose of this article is to describe how this service was instigated and developed. Veterinary advice was to be given not only on immediate medical and surgical problems but also on an overall breeding policy and the control of inherited conditions. The main aim was to keep a healthy group of animals available for fieldwork. The fact that these animals were isolated geographically from other dogs and thus from other canine infectious diseases provided an additional source of interest.

The serendipitous factor that provided veterinary interest in the Antarctic in the first place will be described followed by an account of the work undertaken by each of the three veterinary recruits. Final conclusions will be drawn concerning the goals achieved.

HOW INTEREST AROSE IN VETERINARY PROBLEMS IN THE ANTARCTIC HUSKY

Veterinary presence in the Antarctic arose purely by chance. In 1957, I was in the middle of completing a PhD thesis, on neurological aspects of scrapie in sheep, and needed additional financial support to complete the work. My supervisor was Professor L.P. Pugh, the first professor of Veterinary Clinical Studies at the Cambridge Veterinary School, who approached the Leverhulme Trust on my behalf. An interview was arranged in May of that year with the Trust representative, Sir Raymond Priestley.

At the time I had no reason to be aware that Sir Raymond had been a member of Scott’s British Antarctic Expedition (1910-1913). Although he did not accompany Scott to the South Pole, he was a geologist in the ‘Northern Party’ which planned to carry out an expedition to the west coast, south of Cape Adare. Once the party had been landed, gales and ice prevented it from being recovered by ship, the Terra Nova, and the 6 members had to winter on ‘Inexpressible Island’ on very meagre rations. They managed to build a large igloo and produced oil for lamps and a cooking stove from seal blubber. They survived on strict rations of biscuit and a daily stew of pemmican, seal and penguin meat, known as hoosh. The party, in summer clothing, experienced extreme deprivation over a period of 7 months. Their condition was not only aggravated by cold but also by almost continuous gale (Priestley, 1914).

At the base it was thought they could not survive. There was no radio contact.  Miraculously the party returned to Cape Evans in an emaciated condition, after a 200 mile sledge journey. Unlike Scott, they had been able to prevent the onset of scurvy by eating seal and penguin meat, providing adequate levels of Vitamin C.

Returning now to my interview with Sir Raymond. This was carried out in a relaxed fashion and towards the end the conversation happened to turn on medical problems occurring in huskies in the Antarctic. Apparently a number of dogs were going off their legs, possibly a form of partial paralysis, and had to be euthanised. I suggested that it would be useful to employ a veterinary surgeon to investigate this so-called neurological condition and to give advice on any other veterinary problems. Sir Raymond thought this a good idea but suggested there might be difficulties in gaining official support from BAS, but he would try.

In a letter from Sir Raymond, dated May 1957, he reported that in a cable from the Falkland Islands:-

‘FIDS 105/10/3 HAVE RECEIVED LETTER FROM PALMER CAMBRIDGE UNIVERSITY VETERINARY SCHOOL OFFERING ASSISTANCE AND INDICATING INTEREST IN NERVOUS DISEASES SOMETIMES AFFECTING OUR DOGS STOP CONSIDER MUCH OF VALUE IN LIAISON WITH THE ABOVE DEPARTMENT STOP HAVE ARRANGED FOR PRESERVATION SPECIMENS AND EXTRA COPIES OF VET REPORTS FOR PALMER STOP IN ABSENCE MAIL OPPORTUNITY GLAD IF YOU COULD INFORM HIM AND ESTABLISH NECESSARY LIAISON STOP FACIAL ABSCESSES AND SOME MORTALITY SEE DOG REPORTS TO WHICH SUGGEST HE BE ALLOWED ACCESS = SECFIDS/CKD’   

Another meeting was arranged in Cambridge on 20 May and next day a further telegram arrived from FIDS requesting advice about the composition of a small box of veterinary supplies, to be held at bases and also for information about anaesthetics to be used on huskies. With the help of Dr. L.W. Hall, I put together such a report which was acknowledged by Priestley in June 1957.

Further interest was expressed in a letter from the Falkland Islands Dependencies Scientific Bureau, dated 15 July 1957 requesting a small handbook to be included in the box of veterinary supplies.

I therefore wrote a manual for the veterinary care of huskies, to be carried out by residential medical officers, in the absence of qualified veterinarians. In some 30 pages of quarto, topics included the techniques of intravenous injection, anaesthesia, treatment of wounds, fractures, abdominal surgery and faecal impaction. Comments also referred to the nature of internal and external parasites, care of feet, nephritis, pneumonia, enteritis and post mortem techniques including the selection of tissues and their fixation in 10% formalin.

The manual was completed by September ’57, and this, together with the supplies, was sent down by sea on the next expedition. However, I understand that they never reached their destination. The ship encountered heavy seas and the top hamper had to be relieved by ditching the deck cargo, in which the supplies were loaded – or perhaps they were simply washed overboard (Bellars, personal communication, 1968).

Apart from paraplegia, the huskies suffered from another problem, ‘Signy X disease’, described in a letter from Sir Vivian Fuchs, 8 June 1960. Animals showed inappetance, pyrexia, inflammation of the submandibular lymph glands, abscessation and inflammation of the genitalia. Within 3 days the dogs had either recovered or were dead. In all cases, the animals had been fed raw elephant seal and I suggested that the condition could be bacterial infection, probably caused by Erysipelas rhusiopathiaewhich is known to be widespread in the seal population. The disease occurred on Signy Island, hence its name. Later it was referred to as ‘cervical lymphadenitis’.

I again suggested that a veterinary surgeon should be sent out to the bases to attend to the animals. This idea was favourably entertained by Sir Raymond in his letter (16 Oct 57) although ‘this would be an entirely new departure’. As a result of subsequent deliberations with the Scientific Advisory Committee of FIDS, a Cambridge veterinary graduate, Mike Godsal, was included in the party for the 1963-64 season. His work proved so successful that two more Cambridge graduates Andrew Bellars and Robert Bostelmann were sent south for the seasons 1967-68 and 1973-74.

All huskies, subsequent to the original importation, were bred on the bases, apart from the introduction of a few animals from Greenland and Canada (Bellars, 1969). In the Antarctic, sledge dogs were used for transport of research expeditions as well as for leisure activities. Animals bred on site usually started work when they were a year old and retired at 8 years old. (Bellars, 1969). On retirement an animal may have pulled over 13,000 miles, some did over 19,000 miles (Bostelmann, 1976).

Before the arrival of Godsal, ‘veterinary attention’ had been provided by resident medical doctors, physiologists and dog handlers. As has been mentioned, problems had arisen with so-called Signy X disease and a form of ‘paralysis’, the latter affecting older animals. These conditions clearly needed investigation. But there was also potential interest in studying disease patterns which might arise in a community of dogs, far removed from exposure to the normal range of infectious canine diseases.

M.F. GODSAL

Mike Godsal graduated from the Cambridge Veterinary School in 1963 and interrupted his subsequent appointment as House Surgeon to go to the Antarctic. He visited all the permanently open bases in Grahamland in the summer 1963-64. He saw all the huskies except those in Halley Bay (personal communication, 1964).

He arrived in the Falkland Islands at the end of October 1963. As the ship was due to go on to South Georgia where there was no need of veterinary attention, he offered to stay in the Islands until his departure to the Antarctic. At the time there was no resident veterinary surgeon in the Falklands. There he visited 4 Farm Settlements and discussed various problems with the managers. A common practice was to slaughter sheep in the ‘killing shed’ on the coast. Offal was thrown on to the beach where it could be scavenged by birds, dogs, pigs and even children. The main slaughter house outside Port Stanley had no running water or drainage. All the farm managers admitted seeing ‘cysts’ in sheep offal  Of the 12 pairs of lungs and livers that Godsal examined, 11 contained hydatid cysts.

Four months later when Godsal returned from the Antarctic to Port Stanley he visited more sheep stations and became increasingly concerned about Hydatidosis. When dosing dogs with Tenoban, he was rewarded with copious quantities of tape worms which appeared to be echinococcus. The local medical officer appeared unaware of the dangers despite one young girl having been sent to South America for treatment. Godsal then wrote a letter describing the Hydatid problem in simple terms for distribution to all farmers in the Islands.

When working in the Antarctic, Godsal found most of the huskies to be in good condition. Some of the older ones were not fit for working but had been retained so that they could be examined clinically by him before euthanasia. Some 35 were put down and post mortem examinations carried out. Necropsy samples were fixed in 10% formalin and taken back to the Veterinary School in Cambridge. 

Godsal concluded that nutrition of the huskies was adequate (Nutricon and seal meat) although insufficient quantities were given to prevent weight loss during periods of heavy work (Godsal, 1964). Post mortem findings were disturbing. Some 23 out of 36 animals were found to be suffering from osteoarthritis of the hip and shoulder joints, varying in intensity and distribution. Other abnormalities included prolapsed intervertebral discs, corneal opacities and corneal ulcers. Godsal attributed the corneal ulceration to contamination by volcanic ash.

Godsal treated the huskies for a number of conditions including cauterisation of corneal ulcers with silver nitrate and surgical correction of entropion. While returning on R.R.S. John Biscoe he surgically corrected an intestinal obstruction (seal bones) in a dog called ‘Col’, with the assistance of 2 medical colleagues. He also noted that owing to the generosity of pharmaceutical firms, most of the Antarctic bases were equipped with supplies of acetylpromazine. He advocated its use as a sedative to control fractious huskies when carrying out treatments such as suturing wounds around the eyes. Godsal also contributed to research on occupational osteoarthritis (Bellars and Godsal, 1969).

During the summer of 1965, Godsal had occasion to be in Montreal and flew to the American Air Force Base at Goose Bay (personal communication). There, with the help of the veterinarian in charge of food and hygiene, he obtained 3 huskies for euthanasia from Indian or Eskimo settlements. These dogs had been used for sledging, collecting firewood and occasional hunting trips. Of these three dogs, two were affected with osteoarthritis similar to the cases in the Antarctic. This suggested that neither genetic nor nutritional factors were significant in the aetiology of osteoarthritis in Antarctic huskies.

A.R.M. BELLARS

Three years lapsed before Andrew Bellars was invited to join a BAS expedition during the austral summer 1967 – 68.  His agenda included providing medical and surgical services to the dogs on the bases and continuing Godsal’s survey of diseases and conditions. He was also concerned to find out whether congenital hip dysplasia was a factor which might predispose the animals to develop osteoarthritis later in life.

For this purpose, Bellars borrowed a portable Watson MX2 x-ray from the Cambridge Veterinary School, arranged transport to the base at Stonington where it was installed in the generator shed. There he undertook x-ray examination of the huskies. It should be added that this machine was eventually returned to the Veterinary School, undamaged. By way of provision, Bellars also invited over 30 pharmaceutical companies and veterinary supply firms to donate a comprehensive range of drugs and equipment for work in the Falkland Islands and BAS bases (Bellars, 1969). This dependency on charity seemed to become a feature of veterinary work in the Antarctic.

Bellars travelled out on the R.R.S. John Biscoe, but before reaching the bases, interrupted his voyage at Port Stanley, in the Falkland Islands, where he acted as a temporary resident veterinary surgeon, visiting farms and giving advice over the local radio.

Thereafter he travelled on the John Biscoe and visited all the manned BAS bases and examined all the dogs except for three teams in the field, two at Halley Bay and one at Stonington. In all he examined a total of 203 dogs, recording their medical abnormalities. He x-rayed the pelvis and hips of 25 adult animals and 2 pups as well as a taking a number of lateral radiographs of the spine (Bellars and Godsal, 1969). Using available records he assessed the duration of working life limited by arthritis, whether confirmed or assumed on the basis of abnormal gait or behaviour. He also took slow-motion cine records of the gait of working teams of sledge dogs, from the side, front and rear aspects to see whether abnormal movement might predispose to erosions of hip and shoulder articular cartilages, as previously recognised by Godsal.

He found similar current pathological conditions among the huskies as described by Godsal, including entropion, osteoarthritis, lacerations and bite wounds. He also recorded a brown discolouration of the crowns of teeth in pups, recurrent and intermittent haemorrhages, corneal ulcers, evidence of previous endometritis and one case of anal adenoma. He did not see any cases of Signy X disease, later described as cervical lymphadenitis (Bellars, 1969).

Bellars concentrated his further investigations into three conditions, namely osteoarthritis, entropion and intermittent haemorrhage (later shown to be haemophilia).

OSTEOARTHRITIS

Dogs selected for x-radiography of the hips and pelvis included groups aged less than 18 months, others over 6 years old and some of intermediate age showing signs of premature ‘ageing’. No convincing evidence of hip dysplasia was found in the younger dogs (Bellars and Godsal, 1969). Several of the older animals showed slight abnormalities compatible with erosions of the articular cartilage of the anterior aspect of the femoral head which was usually bilateral. There was also evidence of reshaping of the hip joint and osteophyte formation

Analysis of the patterns of gait, obtained from slow-motion cine films showed that when pulling a sledge both fore and hind-limbs were abducted excessively during protraction, producing a rolling kind of gait with the spine braced horizontally.

In view of the functional, radiographic and pathological evidence, Bellars and Godsal (1969) concluded that the occupational arthritis in the working husky was not related to congenital hip dysplasia but stemmed from mechanical injury to the hip and shoulder joints resulting from pulling heavily loaded sledges. They suggested that the abnormal posture of the dogs caused incorrect distribution of great pressure on the joints leading to malfunction. Bellars and Godsal wondered whether similar work-induced changes occurred in other draught animals.

HAEMOPHILIA    

In total, three dogs were affected, from two consecutive litters, by the same parents. In the second litter, one male animal started bleeding from the mouth when 5 months old, but died within 2 days. Its litter mate, also a dog, started bleeding from the nostril and was euthanised 10 days later. No significant post mortem abnormalities were found. Neither of these dogs was seen by Bellars.

However, the third dog (from the first litter) was examined by him when it was 17 months old. This dog called ‘Malig’ was reported to have started bleeding from one ear when 12 months old. This lasted for 13 days. Over the subsequent years there were 4 more episodes and ‘Malig’ collapsed on one occasion when sledging. Station members were unwilling to use it in a team.

In view of the importance of reaching a definite diagnosis, arrangements were made to transport ‘Malig’ back to England by ship. Quarantine was provided by the Animal Health Trust, at the Small Animal Centre near Newmarket. Laboratory investigations showed that the animal was suffering from true haemophilia.

Bellars completed a large body of work in the Antarctic. He concluded (with Godsal) that the occupational arthritis arose from mechanically-induced forces and was not related to a predisposition to inherited hip dysplasia. However, the occurrence of entropion and haemophilia, both inherited conditions, indicated the importance of planning a breeding programme. To this end, both he and George McLeod drew up a genealogical chart of all 700 animals that had been used in the Antarctic.

Bellars did not revisit the Antarctic but in 1969 went to Greenland where he selected 6 huskies to be sent to BAS bases to improve the breeding stock (personal communication). Through the courtesy of the Royal Greenland Trade Co., he bought the 6 males at Jakobhavn, West Greenland. Unfortunately one of 2 siblings died of heatstroke in Montevideo Harbour.

R.W. BOSTELMANN

The third Cambridge veterinary graduate to visit the Antarctic was Bob Bostelmann who went down during 1972-73. His visit is documented in a paper published in an internal B.A.S report, written in collaboration with N. Macallister and also in a paper published in the Polar Record (1976a) entitled ‘The Management of Sledge Dogs in the Antarctic’. He also wrote two basic manuals for the care of huskies and their pups (‘Kurahound’ and ‘Growapup’).

His voyage to Halley Bay was interrupted by the need to deliver building supplies to Signy Island in the South Orkneys for a boat slipway. On returning to Port Stanley in the Falkland Islands, Bostelmann undertook veterinary work, flying between Camps (farms) and he also gave advice to the Government. He noted that the Hydatid control scheme started in 1964 had resulted in 4 people being sent to Argentina for treatment. He also reported that a number of cats were suffering from a painful paralysis, but did not have the opportunity to examine them.

At the B.A.S base at Halley Bay, Bostelmann had the invidious task of putting down 15 huskies. Clearly this was a thankless business. He carried it out at night at -20C, a cull he described as ‘wrapped up in emotions’.

In his report for B.A.S., Bostelmann gave an account of all registered dogs, the constitution of individual sledge teams and the distances they had travelled. He himself drove his team 1897 miles over a period of 222 days.

It appears that the amount of travelling done with the huskies was restricted at the time by instructions from London. When in the field the dogs lost weight but this was usually regained within three weeks of reaching base. One of the hazards of feeding seal meat was the risk of tracheal cartilage becoming fixed around the base of the dog’s tongue. This could be removed with scissors, usually without anaesthetic.

Bostelmann documented the causes of death. They included exposure, multiple abscessation, strangulation, fighting, abdominal haemorrhage, pyometra, perforated gastric ulcer, fits, lurky (see later), fracture and a case of uterine tumour. One dog had haematuria and another a dilated salivary duct. Others suffered from a ruptured cruciate ligament, intussusception of the small intestine as well as a large number of cases of skin conditions.

Bostelmann’s main work concerned investigating an alternative diet to the traditional nutrican in which the protein was whale meat. As the importation of whalemeat into the UK had been banned, another diet was made up in which soya was the sole protein. A comparison was to be made between this diet and New Zealand nutrican in which the protein was derived from wild deer. Half of a dog team was fed the new diet and the other half given the New Zealand nutrican. After 8 weeks the diets to each group was swapped. Samples of faeces were collected from selected animals and they were weighed. Based on the lack of weight loss and the absorption of fat it was concluded that the soya nutrican was more efficient than the New Zealand version.

Another purpose of Bostelmann’s visit was to investigate ‘foot lurk’. ‘Lurky’ is a term which in B.A.S nomenclature was used in two different senses. A lurky animal describes a nervous animal, difficult to manage and virtually impossible to catch once it got loose. Foot lurk, on the other hand, was a skin condition described initially by Neil Macallister as a ‘pinkish discolouration and swelling on part or all of the pad’. It could extend above the pad and be associated with ‘weeping’. Originally footlurk occurred in three different bases, 200 miles apart. The common factor was that some 15 days earlier the affected dogs had been hauling seal from the beach. A sample subsequently examined in the U.K. showed hairs containing septate hyphae and chains of arthrospores which were very similar in appearance to those of Trichophyton species. There were also micro-abscesses associated with infected hairs (Bostelmann, 1976b). When in the Antarctic, Bostelmann set up a small laboratory (Fig. 1) in an attempt to isolate the organisms from various seal and husky sources, but failed.

The dog breeding programme during Bostelmann’s visit (1972-1973) was extensive but disappointing in that many pups died soon after birth. The average litter size was 6 but pups were usually lost before they had been fed. Early mortality was attributed to overcrowding in the whelping pens and this in turn was due to the tendency of all the bitches coming on heat at the same time (sympathetic oestrus). Some viable pups later turned out to be ‘man shy’ and nervous probably due to the fact that they had not been handled early enough in life.

Bostelmann recorded the history, origin, breeding and training of sledge dogs in the Antarctic in his publication of 1976a. In the same paper he described his own experience on sledging expeditions, the methods of harnessing the dogs to the trace, methods of control and vocal commands. Dogs could be worked for 8 hours with only one rest of 30 minutes. Distances of up to 80km were sledged in a day, but loads of more than 115kg per dog proved difficult to haul. Heavy loads of dog food were obviated by air lifting supplies to strategic posts.

The final outcome of Bostelmann’s work was disappointing for him, in that it was subsequently decided that the use of huskies in the Antarctic was to be phased out, to be replaced by mechanical Skidoos.

In 1963 a message had been sent from London indicating that with increasing use of mechanical aids to travel by B.A.S. it had been decided that no more breeding of dogs was to take place on bases except at Halley Bay where three teams were to be maintained indefinitely (Walton and Atkinson, 1996).

Walton and Atkinson also recorded that in 1991, under pressure from the environmentalists, a new clause was incorporated in the Antarctic Treaty under the title ‘Introduction of non-native species, parasites and diseases’– ‘Dogs shall not be introduced onto land or ice shelves and dogs currently in these areas shall be removed by April 1, 1994’. Effectively this pre-empted the end of an era and of a veterinary presence in the Antarctic.

Apart from a few huskies which over the years had been transported to the U.K. as pets and also some that were exhibited at the Festival of Britain in the Dome of Discovery in 1953, the majority of animals were finally put down and buried. Many years later their bones began to reappear probably because of climate change (C. Rapley, personal communication, 2013).

As a final recognition of the contribution huskies had made to Antarctic travel, 13 of the remaining animals were transported back to their original source, the Inuit village at Inukjuak in northern Canada. It was hoped that these dogs would form a breeding nucleus for the production of future animals.

The operation was called ‘Home for the Huskies’ and was instigated and carried out by John Sweeney (Sweeney, 1994). The thirteen animals were airlifted in a DASH 7 plane on 22 Feb 1994 to Port Stanley. Two days later they were transported to the Cargo Terminal at Heathrow, to be taken by a British Airways 747 across the Atlantic to Boston from where they were driven to Hudson Bay in Canada. From there, Sweeney and two Americans drove 7 of the dogs 300 miles overland to their new owners in the Inuit village. The remaining 6 dogs were flown up.

Unfortunately, the animals did not survive for long, probably succumbing to local infections, despite previous vaccination. 

CONCLUSIONS

What were the benefits of a veterinary presence in the Antarctic? Before its arrival, ‘veterinary care’ had been provided by a resident physician or zoologist, e.g. Drs Neil Orr and Julian Taylor and by dog handlers. Clearly professional help by veterinarians was preferable, especially for the welfare of the animals.

Apart from the medical care of the dogs, the discovery of entropion and haemolytic disease promoted an extensive investigation of the genealogy of the sledge dogs and recommendation of the subsequent breeding. This was later extended into an enquiry into the early deaths of young pups and recommendations for whelping and neonatal care.

A major discovery concerned debilitating arthritis, which affected many of the older animals. Originally, the signs of this condition had been thought to be neurological in origin. Evidence showed that the signs of arthritis did not arise as a consequence of hip dysplasia but was most likely to be mechanical in origin. When pulling heavy loads, considerable force is induced throughout the skeletal system, especially at joint surfaces. Pressure may erode the articular surfaces. On the other hand, there were two diagnostic disappointments in that outbreaks of Signy X disease and pedal ‘lurky’ did not occur during the time of the veterinary tenure. Both conditions were associated with the transport of seal meat, which might suggest the dogs picked up some infectious agent.

My personal contribution was limited to the fortuitous suggestion that a veterinary presence in the Antarctic might be helpful, an idea readily supported by Sir Raymond Priestley. Otherwise I acted as anchor man and supervisor, located in a comfortable Cambridge laboratory, well away from the challenges and discomforts of Antarctic conditions. The Cambridge Veterinary School provided pathological diagnostic facilities and many other services including office space for preparing written reports, all at no cost. One wonders whether such free support would be available in today’s financial climate.

Finally, I was especially pleased to learn that not only was their work appreciated by B.A.S., but the presence of the individual vets on the bases was much appreciated.

REFERENCES

BELLARS, A.R.M. (1969), ‘Veterinary studies on the British Antarctic Survey’s sledge dogs: I. Survey of disease and accidents’. British Antarctic Survey Bulletin, 21, p. 1.

BELLARS, A.R.M. (1969), ‘Hereditary disease in British Antarctic sledge dogs’. Veterinary Record, p. 600.

BELLARS, A.R.M. (1971), ‘Genetic defects in Antarctic dogs’. Journal of Small Animal Practice, 12, p. 493.

BELLARS, A.R.M & GODSAL, M.F. (1969), ‘Veterinary studies on the British Antarctic Survey’ssledge dogs: II. Occupational osteoarthritis’.  British Antarctic Survey Bulletin, 22, p. 15.

BOSTELMANN, R.W. (1976a) ‘The management of sledge dogs in the Antarctic’. Polar Record, 18, p. 25.

BOSTELMANN, R.W. (1976b), ‘Trichophyton infection in Antarctic sledge dogs’. Veterinary Record, 98, p. 425.

BOSTELMANN, R.W. & MACALLISTER, N. (1973), British Antarctic Survey, dog report, reference V/1973/E.

PRIESTLEY, R.E. (1914), Antarctic adventure. Scott’s Northern Party. London: Hurst & Co. Ltd.

SWEENY, J. (1993), Report on the Resettlement of the British Antarctic Survey Huskies.Reproducedcourtesy of the British Antarctic Survey Archives Service ref. SA6/1993/V2

WALTON, K. & ATKINSON, R. (1996), Of dogs and men. Fifty years in the Antarctic. Malvern: Images Publishing (Malvern) Ltd.

ACKNOWLEDGEMENTS

I am very grateful to M.F. Godsal, A.R.M. Bellars and R.W. Bostelmann for their happy participation in this Antarctic venture and for their help in the preparation of this article.

A.C. (Tony) Palmer, MA, ScD, FRCVS, School of Veterinary Medicine, Cambridge