BACTERIOLOGICAL NOMENCLATURE AND TAXONOMY Page 71 CONFERENCE ON VIRUS AND RICKETTSIAL CLASSIFICATION AND NOMENCLATURE New York Academy of Sciences, Section of Biology January 11 and 12, 1952 A Resum6 R . E . Buchanan, Iowa State College, Ames, Iowa The statement approved by the plenary s e s s i o n of the first International Congress of Microbiology a t P a r i s i n 1930 h a s historical significance: "It i s c l e a r l y r e c o g n i z e d t h a t t h e l i v i n g f o r m s w i t h which the microbiologists concern themselves a r e in part plants, in part animals, and in part primitive." The need for adequate considerationof t h e p r o b l e m s of a l l phases of m i crobiological nomenclature was there recognized and emphasized. L a t e r when the International Bacteriological Code of Nomenclature was approved a t the fourth Congress a t Copenhagen, the following wording was authorized: IlB a c t e r i o 1o g i c a 1 n o m e n c 1a t u r e c o n s i d e r s b a c t e r i a , related o r g a n i sm s, and the viruses". - The nomenclature and taxonomy of the v i r u s e s have recently received i n c r e a s e d attention. The plant pathologists interested i n plant v i r u s e s have considered s e v e r a l s y s t e m s of nomenclature proposed. The bacterial virologists have made tentative use of quite different s y s t e m s . The animal virologists in many c a s e s have felt that the time was not opportune for any definite action looking toward either classification o r nomenclat u r e of the animal viruses. It has become increasinglyapparent that the t h r e e groups of virologists, those i n t e r e s t e d i n the higher animals, in the higher plants, and in the bacteriahave much i n common. However, t h e r e was little opportunity for these groups to m e e t to discuss their common problems. The f i r s t major effort to coordinate thinking i n the nomenclat u r e was that of Holmes (1) who in1939 in his Handbook of the Phytopathogenic Viruses proposed that a l l the v i r u s e s be placedin a s e p a r a t e group, the V i r a , and that the Linnaean s y s t e m of binomial nomenclature be applied. The classification was considerably expanded i n Supplement 2 in Bergey's Manual of Determinative Bacteriology ( e d . 6, 1948 ). All v i r u s e s w e r e grouped i n a single o r d e r , the V i r a1 e 8 , with t h r e e subo r d e r s for the bacterial v i r u s e s , the plant v i r u s e s and the animal v i r u s e s respectively. A relatively elaborate classification of each suborder w a s developed, c a r r i e d through to families, g e n e r a and species. Holmes' e a r l i e r classification (1939) of the plant v i r u s e s had considerable acceptance among plant pathologists and the binomial s y s t e m was used by many authors of p a p e r s . However, some of those working i n the field of the animal v i r u s e s felt that the t r e a t m e n t of these o r g a n i s m s by Holmee in Downloaded from www.microbiologyresearch.org by IP: 88.99.165.207 On: Sat, 17 Jun 2017 18:40:51 P a g e 72 BACTERIOLOGICAL NOMENCLATURE AND TAXONOMY 1948 was p r e m a t u r e , and that g r e a t c a r e needed to be used in preventing the straight-jacketing of the nomenclature in this group. It was suggested that s o m e method shouldbe worked out for withholding the final acceptance of all names until there had been an opportunity for the specialists carefully to review the field and to propose a system of nomenclature that would f r o m the beginning have a high degree of acceptability and s tab ility . This widespread feeling on the p a r t df animal virologists led t o the organization through the International Committee on Bacteriological Nomenclature of a special Subcommittee on Viruses to m e e t with the fifth International Microbiological Congress a t Rio de Janeiro in 1950. Dr. C. H. Andrewee of London was designatedas chairman, and asked to s e l e c t the m e m b e r s of such a committee. This he did, and the Committee held s e v e r a l meetings at Rio de J a n e i r o and made a s e r i e s of recommendations to the International Committee on Bacteriological Nomenclature. These r e p o r t s and recommendations have been summarized in previous i s s u e s of this BULLETIN ( 2 , 3, 4). The final outcome of the Rio de Jan e i r o Congress with its attendant conferences was the enlargement of the work of Dr. Andrewes' Subcommittee on Viruses t o m a k e i t representative of all of the major fields of virology. The Committee was asked to o r g a nize as a general Committee onNomenclature and Taxonomy of t h e v i r u s e 8 , with three sub-committees o r Commissions representing each of the three subdivisions of the subject. Dr. Andrewes remained c h a i r m a n of the Subcommittee on Virology andof the Commission on Animal V i r o logy. Later Dr. L. M. Black of Brooklyn was designated a s c h a i r m a n of the Commission on P l a n t Viruses and Dr. J. Craigie of London ar chairman of the Commission on Bacterial Viruses. Lack of opportunity in the p a s t f o r the s e v e r a l groups of virologists to meet to discuss their common p r o b l e m s led the New York Academy of Sciences through i t s Section of Biology to provide f o r a n International Conference on Virus and Rickettsia1 Classification andNomenclature, held in New York January 11 and 12, 1952. The conference was truly international in that many countries w e r e repreeentedandinclusive i n that those interested in the m o s t v a r i e d groups of viruses w e r e in attendance. Abstracts of about thirty p a p e r s and a complete copy of the a d d r e s s of the chairman S i r MacFarlane Burnet were i s s u e d a s a preprint. Several papers of general significance w e r e followed by discussions of the problems of classification of various groups, including the r i c k e t t s i a s , the v i r u s e s o f the higher plants, of insects, of the bacteria, and of higher animals, including such subgroups a s the psittacosis, pox, influenza, arthropod borne , Coxsackie , poliomyelitis and encephalomyocarditis viru s e s . Downloaded from www.microbiologyresearch.org by IP: 88.99.165.207 On: Sat, 17 Jun 2017 18:40:51 BACTERIOLOGICAL NOMENCLATURE AND T A X O N O M Y Page 73 Several of the presentations had to do with general problems of nomenclature of classification and of speciation. D r . E r n s t Mayr of the Museum of Natural History in New York attemptedto shed some light on the p r o blems of taxonomy and nomenclature in the field of microbiology by consideration of species and taxon concepts developed in recent y e a r s in zoology, particularly in ornithology. His overemphasis upon the univer sality of sexual reproductionamong living things, with the inference that r e a l progress in systematic microbiology and virology can be achieved only when something i s known of sexual recombinations i n lower forms such a s the bacteria and the viruses, left the listener with the distinct impression that the application of the standard rules of nomenclature to microorganisms not showing sexual reproduction was scarcely feasible. The suggestion was made that i t mightbe well i f the microbiologists and the virologists left the naming of their organisms to national o r international committees. D r . Burnet emphasized that the valuable developments i n modern population genetics which have led to a better understanding of development and differentiation of species in the higher f o r m s a r e quite without application in the g r e a t groups of agamic microorganisms. He formulated useful c r i t e r i a to be used in the differentiation and classification of the viruses, and emphasized that the organisms cultivated in the laboratory often a r e quite different in certain characters f r o m the organisms a s found in the original host. With some proposed modifications he felt that the outlines of procedure formulated at the Rio de Janeiro Congress could well serve as guides. Dr. Holmes' paper on the problems of nomenclature and classification emphasized some of the special difficulties that a r i s e when virologists in the several fields of bacterial, plant and animal viruses attempt to compose their differences. Holmes, of course, believes that with suitable modifications the Linnaean binomial system of nomenclature is 8 8 tisfactorily applicable to the viruses. In a well-organizedpaper Dr. Andrewes (London) discussed the decisions reached by the Fifth International Microbiological Congress, and outlined the organization that had been evolved to insure orderly p r o g r e s s in the development of a report on nomenclature and taxonomy for presentation a t the sixth Congress a t Rome. It is apparent that Dr. Andrewes and the Committee on Virus Nomenclature and Taxonomy which he heads have a difficult task. Specifically Dr. Andrewes as chairman of a Commission to study one group of animal viruses has the task of organizing a r e p o r t relative to its delimitation, subdivisions and nomenclature, further he has the task af coordinating the work of the s e v e r a l commissions of the animal virology subcommittee, and in turn attempting to formulate r e solutions and suggestions that will be accepted by the plant virologists, the bacterial virologists and the animal virologis<s. Much i n t e r e s t centeredin the rather lively controversy as to whether the Downloaded from www.microbiologyresearch.org by IP: 88.99.165.207 On: Sat, 17 Jun 2017 18:40:51 Page 74 INTERNATIONAL BULLETIN Linnaean system of binomial nomenclature should either now o r eventually be used in the naming of the viruses. Dr. Johnson of Wisconsin insisted that a numbering system for plant viruses of the type proposed by his committee a t the -9msterdam Botanical Congress was preferable. Dr. 3awden of Rothamsted, England, made what appeared to many a s a futile gesture to discredit the binomial system of nomenclature of viruses by ridicule. No poll was taken of the sentiment of the members of the conference a s a whole, but the thinking of certain groups was quite c l e a r . Those interested in the rickettsias seemed reasonably confident that a satisfactory binomial nomenclature in this group was largely developed and increasingly accepted. Similarly there seemed t o be acceptance of binomial nomenclature by those working with the viruses of insects, even though stability has not been achievedas yet. The plant virologists were apparently divided a s to their wishes. Some would develop numbering systems , some felt the binomial system inapplicable and inappropriate, out there was much evidence of increasing acceptance of the Linnaean system in the field of plant pathology. Dr. Andrewes stated that a queetionnaire sent by him to plant virologists in many countries showed opinion to be rather evenly divided for and against the binomial system. One of the American workers pointedout that in a poll of plant virologist. in the United States sentiment i n favor of the binomial system was m a r k edly predominant. No very definite expressionwas given by those working with bacteriophages. The problems in this group seem at present to be largely those of differentiation and classification rather than of nomenclature. Sentiment among the animal virologists is rather more difficult to a s s e s s . There seemed to be no deepseated aversion to the use of the binomial system but it r a s fervently hoped that some method could be devised that would keep animal virology from being engulfed in a nomenclatural mor a s s such a s once threatened nomenclatural stability in bacteriology. Is it not possible in some way to fix uponnames by means of an international agreement? Some were not a v e r s e to Mayr's proposal to have a suitable international committee fix names of taxa by fiat. Several speakers emphasizedthat some groups of viruses a r e much better characterized than a r e others. Some groups maybe readily broken down into genera and speciea, while others a r e still poorly described. A suggestion developed by Dr. Bitancourt of Sao Paulo met with some support. He pointed out a parallelism between the situation existing in the fungi and in the viruses. Scores of genera and thousands of species of fungi cannot be placed with certainty in the formal classification based largely upon the characteristics of the so called perfect stages and a r e allocated to the Fungi Imperfecti, a group created to include those f o r m s whose perfect stage ie unknown or has been completely l o s t from the growth cycle. The Fungi Imperfecti a r e divided into families, genera and apecies and the binomial s y a k m of nomenclature i e applied. Whenever the Downloaded from www.microbiologyresearch.org by IP: 88.99.165.207 On: Sat, 17 Jun 2017 18:40:51 BACTERIOLOGICAL NOMENCLATURE AND TAXONOMY Page 75 perfect stage of one of these l'imperfect fungi" i s discovered i t may be placed in its appropriate genus and if necessary rechristened. It was proposed that a somewhat analogous scheme be adopted for the viruses. Those sufficiently characterizedwouldbe regularly designatedby suitable binomials and the others placedin a group of viruses imperfectly known, and given tentative names with the expectation that when m o r e adequate descriptions a r e available they would be t r a n s f e r r e d to their places in a logical system. Uncertainties a s to classification in some c a s e s could be recognized appropriately by the use of the common Latin phrase indicating uncertainty, there could be viral "G e n e r a I n c e r t a e S e d i s I t as well a s " S p e c i e s I n c e r t a e S e d i s " . This procedure might give a usable nomenclature while m o r e perfect taxonomic outlines were being developed. Several of the papers had special nomenclatural implications. Dr. Black of the Brooklyn Botanical Gardens called attention to the existence of a special problem of classification in the more recently identified group of viruees in which multiplication occurs both in the plant host and in the insect vector. F o u r virus diseases of plants transmitted by leaf hoppers a r e known to belong to this group, Dr. Limaeset of Versailles reviewed i n detail the objections that have been raised to the binomial system of nomenclature as applied to the viruses and concluded that they a r e not rerioue and that some temporary difficultiee might be resolved by use of the grouping "Vira Imperfectar' Dr. Rake of the Squibb Inetitute for Medical Research concluded that c e r tain viruses and rickettsiae a r e sufficiently well characterized to justify their being grouped into genera. The s e v e r a l c r i t e r i a useful i n the development of a classification of the bacterial viruses were outlined by Dr. M a m a of New York University, including serological, morphological physical and physiological characteristics and the results of mixed infections. There a r e a s yet no adequate c r i t e r i a f o r the definition of bacteriophage genera, and the use of binomials may well be postponed until well characterized strains have beenagreed upon as types, and a satisfactory method of maintaining anddistributing such s t r a i n s developed. D r . Coles f r o m hie South African experience presented anincomplete classification of the rickettsias inwhich emphasis was laidupon morphology, serology, culture in eggs, the nature of the m a t r i x etc. He recognized that a satisfactory classification and nomenclature must wait upon m o r e complete understanding of the organisms. Dr. Philip of the Rocky Mountain Laboratory outlined a tentative classification of the R i c k e t t s i a 1 e s in which he recognized one family, three genera and three subgenera. Dr. Bergold of the Canadian Laboratory of Insect Pathology and D r , Steinhaus of the University of Californiaaccepted in principle the binomial system of naming the ineect viruses, but pointed out some of the p r e s e n t difficult problems both of classification and of nomenclature. The f o r m e r ruggested that some international committee might well be given some Downloaded from www.microbiologyresearch.org by IP: 88.99.165.207 On: Sat, 17 Jun 2017 18:40:51 Page 76 INTERNATIONAL BULLETIN degree of authority relative to the names finally to be accepted, a committee that could be consultedby investigators. Dr. Steinhaus listed two genera of insect viruses that seemed to be well established, together with their species , and withdesignationof the type species. Two other genera were includedinthecategoryof G e n e r a e t S p e c i e s I n c e r t a e S e dis. The problems to be solved relative to nomenclature of the s e v e r a l groups of animal viruses vary widely. Dr. Karl Meyer of the University of California posed the difficulties in the psittacosis -ornithosis group of l a r g e viruses and protested the unsatisfactory nomenclature. Dr. Rake of the Squibb Institute, New York, suggested that the lymphogranuloma-psittacosis group showed closer relationships to the rickettsiae than to the v i r u s e s . It i s apparent from the reports given by Dr. F r a n c i s of the U n i versity of Michiganandby S i r MacFarlane Burnet of Melbourne, Australia that substantialprogress i s being made inanunderstanding of the influellza v i r u s group and i n eventual systematization of the nomenclature. Dr. Hammon of the University of Pittsburgh believes that there is insufficient knowledge of the arthropod-borne encephalitis viruses to make possible at present a rational use of the binomial system of nomenclature, and that the classificationproposed by Holmes wae premature. Dr. Lepine of the P a s t e u r Institute, P a r i s a l s o discussed this group and indicated that subrtantial p r o g r e s s is being made. A suggested grouping of the arthropod viruses was noted, emphasizing that the s e v e r a l v i r u s e s appear to belong together although causing very distinct diseases and with distinct a r t h r o pod vectors. A suggested generic diagnosis for the Coxsackie viruses and diagnoses of two groups within the genus was presented by Dr. D a l l dorf of the New York State Department of Health. He was able to differentiate ten types i n the f i r s t group and three i n the second. Dr. Malnick of Yale University pointed out certaindifficulties in the acceptance of the two groups of Coxsackie virus a s proposed by Dalldorf. A constructive c r i t i c i s m of the Holmes's classification of the poliomyelitis viruses was presented by Dr. Rhodes of the University of Toronto. The poliomyelitis group and c r i t e r i a f o r diagnosis of a single genus with two o r three species were discussed by D r . Koprowski of the LederPe Laboratories, emphasis being laid upon morphology as determined by the use of the electron microscope. A schema for the encephalomyocarditis group wae outlined by Dr. Warren of the Army Medical Service Graduate School, Washington. The need of adequate and reliable type culture collections for the viruses was emphasized by several speakers. Dr. Smadel of the Army Medical Service Graduate School outlined the service of the Virus and Rickettsial Registry U. S.A. , and Dr. McKinney the need for an adequate culture collection for the use of those interested in plant virology. The l a s t session of the Conference was devoted to an open discussion of the problems of virus nomenclature under the chairmanship of S i r Mac- Downloaded from www.microbiologyresearch.org by IP: 88.99.165.207 On: Sat, 17 Jun 2017 18:40:51 INTERNATIONAL BULLETIN Page 77 Farlane Burnet. The results of the conference and the papers presented a r e in process of publication by the New York Academy of Science. They will constitute a reference book of r e a l value to a l l i n t e r e s t e d in the virus field and in its systematics. In summary, it may be noted that Dr. Andrewes and his committee and commissions on virological nomenclature have the important task of p r e paring nomenclatural and taxonomic recommendations to be submitted to the next International Congress of Microbiology in September 1953. As f a r as practicable, virologists should take an active i n t e r e s t in the work of this committee and render all needed assistance. The task of the committee is complicated and difficult. Among its s e v e r a l assignments is that of making a definite proposalfor a suitable date of departure for the nomenclature of the viruses. This fixing of a date is needed, no matter what form of nomenclature may be finally proposed or approved. There a r e many possible solutions to the problem. The deliberations of the conference and a survey of the actions that have been taken by international agreement in botany, zoology and bacteriology suggest the general nature of some of the possible solutions. To the reviewer, four s e e m worthy of mention. 1. The Committee might consider recommending that all names validly published in the literature of virology be regarded as available f r o m the standpoint of priority (essentially the practice in m o s t branches of biology 1. 2. The Committee might consider recommending that the starting point in virological nomenclature be some published monograph of the viruses, and all older names nottherein recognizedbe ignored. Such an arrangement is in effect in botany for some groups of the fungi and of the algae, In the event of such a proposal, the monographof Holmes would be among those to be considered. 3. The Committee might consider the advisability of preparing or of supervising the preparation of an authoritative monograph on the viruses, describing all the genera and higher taxa recognized, listing and naming the species accepted, and designating type species and type genera. O r the monograph might be definitive for certain of the better known groups of the viruses, with those not well known o r indefinitely characterized placed in some such category a s Vira imperfecta or Viraincertae sedis. Those organisms indefinitely placed would l a t e r be given recognized s t a tus when better characterized. Such a proposal would essentially be a recommendation for the conservation of a list of generic names with designation of the types. 4. The Committee may well find that the time available before the 1953 Congrese is too limited to complete fully the task assigned. A recommendation that additional time be granted the Committee would doubtless be approved. It is to be hoped, however that substantial p r o g r e s s may have been made before any postponement is requested. There is rapidly Downloaded from www.microbiologyresearch.org by IP: 88.99.165.207 On: Sat, 17 Jun 2017 18:40:51 Page 78 BACTERIOLOGICAL NOMENCLATURE AND TAXONOMY mounting p r e s s u r e , c e r t a i n l y in America, f o r definitive action on v i r u s nomenclature. A s u r v e y of r e c e n t l i t e r a t u r e r e v e a l s that binomial n a m e s f o r the v i r u s e s are increasingly employed and a r e accepted by a l a r g e proportion of the virologists. Obviously no author of a text which t r e a t s of the v i r u s e s , e i t h e r of plants or animals, c a n ignore the n a m e s that a r e comiag into u s e . It s e e m s probable that the flood p r e s s u r e m a y well be held under control until 1953 but it is to be hoped that the r e g u l a t o r y dev i c e s m a y be p r e t t y well completed by that t i m e . Downloaded from www.microbiologyresearch.org by IP: 88.99.165.207 On: Sat, 17 Jun 2017 18:40:51
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