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Professor Peter L. Long: An Appreciation

M. W. Shirley
Institute for Animal Health, Compton Laboratory, Compton, Newbury, Berks. RG 20 7NN, UK, shirley@bbsrc.ac.uk

If asked to think of the most influential coccidiologist of the past four decades, the chances are that the name of Peter Long will feature at, or near, the top of most people's lists. If asked to think of the most influential coccidiologist of the past four decades who is also the most approachable and sociable, the likelihood is that Peter will be at the top of everyone's list! Peter has the rare gift of scientific excellence combined with a warm demeanour and fondness for bon homie that has endeared him to all who have had the pleasure of his acquaintance.
I first became a colleague in 1967 when I started work at the Houghton Poultry Research Station (HPRS) where Peter was a principal scientist looking to recruit a new member of his team. At that time he had not yet gained his PhD but he was established comfortably as a coccidiologist of increasing international renown and was a natural successor to Clifford Horton-Smith who had also previously graced HPRS with distinction and charisma. A symmetry in the three generations was to emerge - Clifford had been a mentor to Peter as Peter was to become to me. Moreover, there was to be further symmetry in the way that the academic career of Peter developed subsequently with how mine was to shape up under his guidance.
Edward Tyzzer was working on his Magnum Opus to be published in 1929 when Peter was born in London in 1928. Peter developed into a good sportsman with a special love of cricket - a game at which he was especially skilled and which he played on-and-off for several decades. He supplemented his cricket with both hockey and squash and his competitive nature at sports was mirrored by his determination to succeed professionally. 
He began his professional life working on poultry parasites at the Central Veterinary Laboratory at Weybridge, Surrey as a laboratory technician with Dr Clifford Horton-Smith. Only four years later, however, he was conscripted into the Royal Army Medical Corps and served in Egypt, Malta and Palestine. 
In 1949, following demobilisation, Peter and Clifford were reunited at the Ashford Hospital, Middlesex and then a more durable union followed with their relocation to the HPRS: Peter becoming the research technician for Horton-Smith within the Parasitology department working on a programme of research directed principally towards coccidiosis of poultry. Without any formal academic qualifications, Peter studied part-time to become a Member of the Institute of Medical Laboratory Technology in 1952 and obtained Fellow status by thesis in 1956.
A steady stream of publications was beginning to appear and a strong desire to publish his data was to define his entire working life. Some early publications included a very useful treatise on E. maxima [Long, P. L. (1959). A study of Eimeria maxima Tyzzer, 1929, A coccidium of the fowl (Gallus gallus). Annals of Tropical Medicine and Parasitology, 53, 325-333] that remains one of the few definitive studies on this major avian species. By now Peter was progressing through the ranks from his original grading of "Assistant Experimental Officer" such that by 1963 he was able to spend a sabbatical year at Cornell University in the USA working with the eminent poultry pathologist, Professor P. P. Levine. This very positive scientific experience left an indelible mark on his subsequent career and much later he was to return to the USA to the University of Georgia where he spent the final 10 years of his active working life.
Peter's taste for working abroad was strengthened in 1963 and 1964 with two visits to the Lebanon, where he worked on poultry coccidiosis and taught parasitology at the Institut de Recherches Agronomiques.
In 1964, and through further part time studies, Peter became a member of the Institute of Biology in the UK and in 1971 Peter was awarded a PhD by Brunel University for his work on the avian coccidia and in 1977 was awarded a DSc. His association with Brunel University led to a strong friendship with Professor Roland J. Terry, who subsequently paid a professional visit to Peter when he was at Athens.

Throughout the 1960's and 1970's Peter maintained a high output of publications from the parasitologists at HPRS with an impressive list of personal papers that laid much of the groundwork for later developments in the control and study of the avian coccidia including:

  • two of the few papers on Eimeria to be published in the journal Nature [Long, P. L. (1965). Development of Eimeria tenella in avian embryos. Nature 208, 509-10 and Long, P. L. (1970c). Development (schizogony) of Eimeria tenella in the liver of chickens treated with corticosteroid. Nature 225, 290-1]

  • pioneering studies on the growth of different species of Eimeria in avian embryos that would eventually help in the development of live attenuated vaccines [Long, P. L. (1966). The growth and attenuation of some species of Eimeria in avian embryos. Parasitology 56, 575-81 and Long, P. L. (1972b). Eimeria tenella: reproduction, pathogenicity and immunogenicity of a strain maintained in chick embryos by serial passage. J Comp Pathol 82, 429-37]

  • the isolation and identification of different species [e.g. Long, P. L. (1973c). Studies on the relationship between Eimeria acervulina and Eimeria mivati. Parasitology 67, 143-55 and Long, P. L. (1974). Experimental infection of chickens with two species of Eimeria isolated from the Malaysian jungle fowl. Parasitology 69, 337-47]

  • studies on immunity, often with Elaine Rose [Long, P. L., and Rose, M. E. (1972). Immunity to coccidiosis: effect of serum antibodies on cell invasion by sporozoites of Eimeria in vitro. Parasitology 65, 437-45

  • studies on chemotherapy [Long, P. L., and Millard, B. J. (1968). Eimeria: effect of meticlorpindol and methyl benzoquate on endogenous stages in the chicken. Exp Parasitol 23, 331-8]

  • the first studies to provide quantitative data on the incidence of Eimeria species in poultry houses [Long, P. L., Tompkins, R. V., and Millard, B. J. (1975). Coccidiosis in broilers: evaluation of infection by the examination of broiler house litter for oocysts. Avian Pathology 4, 287-294].

This was indeed a golden period for Peter and his acclaim grew within HPRS, where he was appointed successor to Dr Donald L. Lee as Head of Parasitology in 1972; nationally, for example, he was awarded the Tom Newman Medal for Poultry Research from the British Poultry Breeders and Hatcheries Association in 1971 and made an honorary member of the British Veterinary Poultry Association in 1972; and internationally where he consolidated his position as arguably the leading scientist working on the biology of avian coccidia.
In 1973 Peter worked with Datus M. Hammond to edit "The coccidia", a book that was to become the first in a series of volumes on the biology of the coccidia and coccidiosis. These books, most especially the 1982 volume entitled "The biology of the coccidia" have stood the test of time and continue to be some of the most used reference texts. The most recent volume, now over 15 years old and entitled "Coccidiosis in man and domestic animals", is a further reflection of Peter's commitment and ability to work with others to effectively deliver knowledge of the avian coccidia into the public domain. 
Within the UK Peter helped to establish the forum that is now known as the European Coccidiosis Discussion Group and he was one of the founding members of what was to become the British Society for Parasitology, which later made him an honorary member. 
In 1979, Peter left the HPRS to succeed Professor W. Malcolm Reid as Professor of Parasitology and Chairman of the Faculty of Parasitology at the University of Georgia. In 1983, Peter was further recognised by the University by the title, which he held until 1990 when he retired back to the UK, of D.W. Brooks Distinguished Professor of Poultry Science.
During his years at HPRS, Peter was most ably supported scientifically and technically by Brian Millard and, if there were an equivalent anywhere else in the world, he most definitely found that person in Joyce Johnson. Peter and Joyce developed a wonderful working rapport and the transition of his science from HPRS to the University of Georgia seemed almost seamless with virtually no interruption to the stream of papers.
Peter's later contributions to coccidiosis research continued to reflect his desire to improve and understand the control of disease and his early work in Georgia was pivotal to the first understandings of the relationship between pathological changes in the intestines and body weight changes of vaccinated or naive chickens after challenge, findings that were to become important considerations in the evaluation of vaccines [e.g. Long, P. L., Johnson, J., and Wyatt, R. D. (1980). Eimeria tenella: clinical effects in partially immune and susceptible chickens. Poult Sci 59, 2221-4]. In addition he pursued work in the USA on the development of precocious, attenuated lines of Eimeria as well as addressing issues of disease control more generally.
In total, Peter published more than 200 scientific papers, countless popular articles and very many book chapters. He also helped stage the Coccidiosis Conference in Georgia in 1985.
The scientific excellence of Peter and his contribution to research on coccidia and coccidiosis has been recognised many times and, in addition to the Tom Newman Award, he was the first recipient of the Gordon Memorial Medal (1983); the Merck Award for Research from the American Poultry Science Association (1984); and the Creative Research Medal from the University of Georgia, Athens.
Throughout his career Peter influenced and mentored many colleagues including several PhD students (about whom he still speaks with great warmth and pride) as well as many more established researchers across the globe. He enjoyed interacting with, or working alongside, such luminaries as Alan Pierce, Akira Arakawa, Stan Ball, Peter Bedrnik, David Chapman, Aggie Fernando, Datus Hammond, Thomas Jeffers, Joyce Johnson, Len Joyner, P. P. Levine Lazlo Pellerdy, Larry McDougald, Malcolm Reid, Elaine Rose, Erich Scholtyseck and Ray Williams, to name just a few.
When David Chapman and I published a recent history of the Houghton strain, a strain that is now so firmly embedded within research into Eimeria spp. as the parasite used for the derivation of a genome sequence, we dedicated the paper to both Peter Long and Elaine Rose. Without the direct inputs of Peter on both the science and the people, the current landscape of coccidiosis research would almost certainly be very different from what we have now and probably less at the cutting-edge of microbiological research. Many of us working on Eimeria parasites might care to reflect upon the words of Isaac Newton to Robert Hooke on 5th. February 1676, that "If I have seen further, it is by standing on the shoulders of giants". Peter Long is a giant in the history of coccidiosis research.

  • A selection of other publications by Peter Long that further illustrates his very significant contribution to studies on the avian coccidia

  • Long, P. L. (1967). Studies on Eimeria mivati in chickens and a comparison with Eimeria acervulina. J Comp Pathol 77, 315-25.

  • Long, P. L., and Millard, B. J. (1967). The effect of Meticlorpindol on Eimeria infections of the fowl. Vet Rec 81, 11-5.

  • Long, P. L. (1968). The effect of breed of chickens on resistance to Eimeria infections. Br Poult Sci 9, 71-8.

  • Long, P. L. (1968). The pathogenic effects of Eimeria praecox and E. acervulina in the chicken. Parasitology 58, 691-700.

  • Long, P. L., and Horton-Smith, C. (1968). Coccidia and coccidiosis in the domestic fowl. Adv Parasitol 6, 313-25.

  • Long, P. L. (1969). Observations on the growth of Eimeria tenella in cultured cells from the parasitized chorioallantoic membranes of the developing chick embryo. Parasitology 59, 757-65.

  • Long, P. L. (1970). Coccidiosis: development of new techniques in coccidiostat evaluation. Exp Parasitol 28, 151-5.

  • Long, P. L. (1970). Eimeria tenella: chemotherapeutic studies in chick embryos with a description of a new method (chorioallantoic membrane foci counts) for evaluating infections. Z Parasitenkd 33, 329-38.

  • Long, P. L. (1970). Some factors affecting the severity of infection with Eimeria tenella in chicken embryos. Parasitology 60, 435-47.

  • Long, P. L. (1970). Studies on the viability of sporozoites of Eimeria tenella. Z Parasitenkd 35, 1-6.

  • Long, P. L., and Rose, M. E. (1970). Extended schizogony of Eimeria mivati in betamethasone-treated chickens. Parasitology 60, 147-55.

  • Long, P. L., and Milne, B. S. (1971). The effect of an interferon inducer on Eimeria maxima in the chicken. Parasitology 62, 295-302.

  • Long, P. L. (1971). Schizogony and gametogony of Eimeria tenella in the liver of chick embryos. J Protozool 18, 17-20.

  • Long, P. L. (1972). Eimeria mivati: reproduction, pathogenicity and immunogenicity of a strain maintained in chick embryos by serial passage. J Comp Pathol 82, 439-45.

  • Long, P. L. (1972). Observations on the oocyst production and viability of Eimeria mivati and E. tenella in the chorioallantois of chicken embryos incubated at different temperatures. Z Parasitenkd 39, 27-37.

  • Long, P. L., Burns Brown, W., and Goodship, G. (1972). The effect of methyl bromide on coccidial oocysts determined under controlled conditions. Vet Rec 90, 562-7.

  • Long, P. L. (1973). Endogenous stages of a 'chick embryo-adapted' strain of Eimeria tenella. Parasitology 66, 55-62. 

  • Long, P. L. (1973). The growth of avian coccidia in the developing chick embryo and in cultured cells . Folia Vet Lat 3, 89-109.

  • Long, P. L., Fernando, M. A., and Remmler, O. (1974). Experimental infections of the domestic fowl with a variant of Eimeria praecox from the Ceylon jungle fowl. Parasitology 69, 1-9.

  • Long, P. L., and Rowell, J. G. (1975). Sampling broiler house litter for coccidial oocysts. Br Poult Sci 16, 583-92.

  • Long, P. L., Millard, B. J., Joyner, L. P., and Norton, C. C. (1976). A guide to laboratory techniques used in the study and diagnosis of avian coccidiosis. Folia Vet Lat 6, 201-17.

  • Long, P. L., and Millard, B. J. (1976). The detection of occult coccidial infections by inoculating chickens with corticosteroid drugs. Z Parasitenkd 48, 287-90.

  • Long, P. L., and Millard, B. J. (1976). Studies on site finding and site specificity of Eimeria praecox, Eimeria maxima and Eimeria acervulina in chickens. Parasitology 73, 327-36.

  • Long, P. L., and Rose, M. E. (1976). Growth of Eimeria tenella in vitro in macrophages from chicken peritoneal exudates. Z Parasitenkd 48, 291-4.

  • Long, P. L., Millard, B. J., and Shirley, M. W. (1977). Strain variations within Eimeria meleagrimitis from the turkey. Parasitology 75, 177-82.

  • Long, P. L., and Millard, B. J. (1978). Coccidiosis in broilers: the effect of monensin and other anticocidial drug treatments on oocyst output. Avian Pathology 7, 373-381.

  • Long, P. L., and Millard, B. J. (1978). Studies on Eimeria grenieri in the guinea fowl (Numida meleagris). Parasitology 76, 1-9.

  • Long, P. L., and Millard, B. J. (1979). Immunological differences in Eimeria maxima: effect of a mixed immunizing inoculum on heterologous challenge. Parasitology 79, 451-7.

  • Long, P. L., and Millard, B. J. (1979). Rejection of Eimeria by foreign hosts. Parasitology 78, 239-47.

  • Long, P. L., and Millard, B. J. (1979). Studies on Eimeria dispersa Tyzzer 1929 in turkeys. Parasitology 78, 41-52.

  • Long, P. L., Millard, B. J., and Lawn, A. M. (1979). An unusual local reaction to an intracellular protozoon parasite Eimeria dispersa. Z Parasitenkd 60, 193-5.

  • Long, P. L. (1970). Anticoccidial drugs: factors affecting pathogenicity of avian coccidia. Exp Parasitol 28, 4-10.

  • Long, P. L., Johnson, J., and Reyna, P. (1980). The use of sentinel birds to monitor coccidial infection in litter pens. Avian Dis 24, 435-45.

  • Long, P. L., Johnson, J., and Wyatt, R. D. (1980). Eimeria tenella: clinical effects in partially immune and susceptible chickens. Poult Sci 59, 2221-4.

  • Long, P. L., Johnson, J., and Wyatt, R. D. (1981). Pathological and clinical effects of Eimeria tenella in partially immune chickens. J Comp Pathol 91, 581-7.

  • Long, P. L., and Jeffers, T. K. (1982). Studies on the stage of action of ionophorous antibiotics against Eimeria. J Parasitol 68, 363-71.

  • Long, P. L., Johnson, J., and Gore, T. C. (1982). Attenuation of a strain of Eimeria mivati of U.S. origin by serial embryo passage. Avian Dis 26, 305-13.

  • Long, P. L. (1984). Gordon memorial lecture. Coccidiosis control: past, present and future. Br Poult Sci 25, 3-18.

  • Long, P. L., and Joyner, L. P. (1984). Problems in the identification of species of Eimeria. J Protozool 31, 535-41.

  • Long, P. L., Johnson, J., and Baxter, S. (1985). Eimeria tenella: relative survival of drug-resistant and drug-sensitive populations in floor pen chickens. Poult Sci 64, 2403-5.

  • Long, P. L., Johnson, J., and McKenzie, M. E. (1988). Anticoccidial activity of combinations of narasin and nicarbazin. Poult Sci 67, 248-52.

 

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