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Serratia Proteamaczdans (Paine and Stansfield) Comb INTERNATIONAL JOURNALOF SYSTEMATIC BACTERIOLOGY, OCt. 1978, p. 503-510 Vol. 28, No. 4 0020-7713/78/0028-0503$02.00/0 Copyright 0 1978 International Association of Microbiological Societies Printed in U.S. A. Serratia proteamaczdans (Paine and Stansfield) comb. nov., a Senior Subjective Synonym of Serratia Ziquefaciens (Grimes and Hennerty) Bascomb et al. PATRICK A. D. GRIMONT,? FRANCINE GRIMONT,? AND MORTIMER P. STARR Department of Bacteriology, University of California, Davis, California 95616 In 1919, Paine and Stansfield attributed a leaf-spot disease of the tropical plant Protea cynaroides to Pseudomonas proteamaculans. This organism, placed at various times in the genera Xanthomonas, Erwinia, and Enterobacter and now known as Erwinia proteamaculans (Paine and Stansfield 1919) Dye 1966, is listed in the first (1976) draft of the Approved List of Bacterial Names. We demonstrate here-on the basis of biochemical properties, polynucleotide se- quence relatedness, and pathobiological capacity-the identity of (i) the only strain (ATCC 19323 = Dye ZL1 = ICPB XP176 = NCPPB 245; here designated as the type strain of P. proteamaculans) of E. proteamaculans still extant from the Paine and Stansfield study with (ii) Serratia liquefaciens biotype Clc. Based on the results of our study, we recommend the transfer of E. proteamaculans to the genus Serratia. Serratia proteamaculans (Paine and Stansfield 1919) comb. nov. is to be regarded as a senior subjective synonym of S. liquefaciens (Grimes and Hennerty 1931) Bascomb et al. 1971. In 1919, Paine and Stansfield (29) observed in bacteria that are associated with plants have Kew Gardens (London) what they believed to been assigned to the genus Erwinia solely on be a leaf-spot disease of the tropical plant Protea the basis of their phytopathogenicity (40).These cynaroides (vernacular name, King Protea). factors led us to undertake a survey on the They attributed the disease to a bacterium relationships between plant and medical enter- which they isolated from the leaf spots and obacteria, in the course of which we found Er- named Pseudomonas proteamaculans (29). Al- winia proteamaculans to be identical with one though no bacterial leaf-spot disease of Protea biotype (Clc) of Serratia liquefaciens. After has been reported since 1921 (28),the taxonomic checking the authenticity of the only extant position of the alleged causative agent has strain of the collection on which Paine and changed several times, as evidenced by the fol- Stansfield based their description of P. protea- lowing list of objective synonyms: Pseudomonas maculans (ATCC 19323 = Dye ZL1 = ICPB proteamaculans Paine and Stansfield 1919 (29); XP176 = NCPPB 245), we demonstrated the Bacterium proteamaculans (Paine and Stans- identity of this strain with S. Ziquefaciens bio- field 1919) Elliott 1930 (16);Phytomonasprotea- type Clc (19) by comparison of biochemical maculans (Paine and Stansfield 1919) Bergey et reactions, polynucleotide wquence relatedness, al. 1930 (4); Xanthomonas proteamaculans and pathobiology upon inoculation of leaves of (Paine and Stansfield 1919) Burkholder 1948 Protea cynaroides. Relevant nomenclatural (11); Erwinia proteamaculans (Paine and changes are proposed. Stansfield 1919) Dye 1969 (14). In the eighth (1974) edition of Bergey’s Manual, Lelliott com- MATERIALS AND METHODS mented upon Erwinia proteamaculans in the Bacterial strains. The histories of the bacterial following manner: “Possibly an Entero bacter strains studied are given in Table 1. Cultures of these sp.; excluded from Erwinia because the only strains were freeze-dried for long-term conservation known extant cotype produces lipwe, and lysine (34) and were also maintained on semisolid yeast extract nutrient agar (19) during this study. and ornithine decarboxylases” (25). Bacteriological methods. Polygalacturonate deg- The taxonomy of plant-pathogenic bacteria radation was studied using the pectate semisolid agar has suffered from “disciplinal insularity” (35) of Starr et al. (36). The caprylate-thallous agar me- and from the curious practice in which most, if dium, selective for members of the genus Serratia, has not all, peritrichous, gram-negative, rod-shaped been described previously elsewhere (37). All other bacteriological tests were performed exactly as de- t Permanent address: Service des Enterobacteries, Institut scribed by Grimont et al. (19). All tests were carried Pasteur, F-75724 Paris, Cedex 15, France. out at 30°C, unless otherwise stated. 503 504 GRIMONT, GRIMONT, AND STARR INT. J. SYST.BACTERIOL. TABLE1. Bacterial strains studied" Strain designation Species Strain historyb ICPB Grimont et al. 119) Xanthomonasproteamaculans ..... XP176 - NCPPB 245 t NCTC 394 t S. G. Paine (29) Serratia Iiquefaciens .............. 3989 508 CCM 412 c CCEB 309; from insect, Saperda carcharias S. liquefaciens .................... 3991 275 Le Minor 5-68 c CDC 6136-66; centrotype of phenon C1 (19) S. liquefaciens .................... 3994 503 ATCC 14460, type strain of S. liquefaciens (19, 33) S. plymuthica .................... 4002 510 CCM 640 +- ATCC 183; type strain of S. ply- muthica (19) S. marcescens .................... 4008 296 Brisou 5751; from water; centrotype of phenon A (19) S. marinorubra ................... 4003 288 Sneath D119 t ZoBell 511; type strain of S. marinorubra (19) Enterobacter aerogenes ........... 2856 A1 CIP 60-86 +- ATCC 13048; neotype of E. aero- genes (20) E. cloacae ....................... 2857 c1 CIP 60-85c- ATCC 13047; neotype of E. cloacae (20) Escherichica coli K-12 ............. 4023 - Strain 55 from N. Datta a In addition to the above-mentioned Serratia strains, the following strains from the study of Grimont et al. (19) were used in plant inoculation experiments: Serratza liquefaciens 221,390, and 509;S. marcescens 5,60,81, 222, 246, 324, 481, and 1264; S. marinorubra 37, 377, and 530; S. plymuthica 34 and 392; Serratia sp. 38 and 10:3. ATCC, American Type Culture Collection, Rockville, Md.; CCEB, Czechoslovak Collection of En tomoge- nous Bacteria, Prague; CCM, Czechoslovak Collection of Microorganisms, Brno; CDC, Center for Disease Control, Atlanta, Ga.; CIP, Collection de l'Institut Pasteur, Paris, France; ICPB, International Collection of Phytopathogenic Bacteria, Davis, Calif; NCPPB, National Collection of Plant Pathogenic Bacteria, Harpenden, England; NCTC, National Collection of Type Cultures, London, England. DNA-DNA hybridization studies. The medium containing only denatured labeled XP176 DNA, and used in the labeling of deoxyribonucleic acid (DNA) a sample containing only native labeled XP176 DNA. with [3H]thymidine consisted of (per liter): KH2P04, After incubation, four 0.24 samples were removed 2 g; K2HPO4,7 g; MgS04 7H20,O.l g; (NIC)&Od, 1g; from each hybridization mixture and transferred to glucose, 1 g; CaSamino Acids, 15 g; and NaCl, 0.5 g. tubes containing 0.8 ml of reaction mixture (0.4 mM Sixty milliliters of sterile medium in a Nett flask was ZnSOr, 0.15 M NaC1,0.03 M sodium acetate buffer at inoculated with strain XP176. When the growth pH 4.8, and 20 pg of sheared and denatured calf thymus reached 40 Klett units, deoxyadenosine and [3H]thy- DNA per ml). Two tubes from each reassociation midine (specific activity, 20 Ci/mmol) were added at mixture were each treated with 5.0 pl (100 U) of S1 final concentrations in the medium of 250 and 8.0 nuclease (Sigma Chemical Co., St. Louis, Mo.) for 20 pg/ml, respectively. The bacterial culture was har- min at 60°C, and two tubes were treated identically vested when it reached 240 Klett units. Radioactive but without the addition of the S1 enzyme. DNA DNA was extracted and purified by a published pro- duplexes remaining after this treatment were precipi- cedure (8). Extraction and purification of unlabeled tated by the addition of 1.0 ml of ice-cold 10% trichlo- DNAs and shearing of both labeled and unlabeled roacetic acid and collected on glass-fiber filters (What- DNAs were done by the method of Brenner et al. (9), man GF/F). Filters were washed with four 2.5-ml except that the purified DNAs were dialyzed against volumes of ice-cold 5% trichloroacetic acid and then 0.042 M NaCl before shearing by sonic oscillation. with 2 ml of acetone. The filters were dried and put For hybridization experiments, the methods of into vials containing 5 ml of scintillant (4 g of Omni- Crosa et al. (12) and of Schiewe et al. (32) were fluor per liter of toluene), and the radioactivity was followed with slight modifications. A mixture of 0.1 pg measured in a Beckman model LS3145P liquid scintil- of 3H-labeled DNA and 150 pg of unlabeled DNA in lation spectrometer. The degree of polynucleotide se- 0.042 M NaCl (total volume, 0.75 ml) was denatured quence homology was calculated by determining the in a boiling-water bath for 4 min and immediately ratio between the average counts in the nuclease- quenched in ice. The NaCl concentration was then treated and nuclease-untreated samples. Results were adjusted to 0.42 M by addition of 0.25 ml of 1.554 M then normalized to the homologous reaction. NaCl. This hybridization mixture was then incubated Plant inoculations. Four pots of Protea cyna- for 16 h in a 60°C water bath. Included as controls roides (vernacular name, King Protea) were obtained with each group of DNA reassociations were a homol- from Tropic World Inc. (Escondido, Calif.), and two ogous DNA reassociation (XP176/XP176), a sample others were obtained from Green Valley Nurseries VOL. 28,1978 SERRATIA PROTEAMACULANS COMB. NOV. 505 (Escondido, Calif.). Overnight cultures of various bac- ual of Determinative Bacteriology (4, 5) cited teria in Trypticase soy broth (BBL Microbiology Sys- this description uncritically, under various ge- tems) were deposited (10 pl) on detached leaves of neric names. In the seventh (1957) edition of Protea cynaroides. Leaves were then lightly stabbed Bergey’s Manual (6), the description was cor- with a sterile platinum wire through the deposited drop of culture and were kept in a humid chamber at rected as follows: “Gram-positive (Paine and 30°C.
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