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THE HYBRID-SUBSTANCE of the ERYTHROCYTES of the HYBRIDS BETWEEN COLUMBA LIVIA and STREPTOPELIA RISORIA' T Has Been Thought

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THE HYBRID-SUBSTANCE of the ERYTHROCYTES of the HYBRIDS BETWEEN COLUMBA LIVIA and STREPTOPELIA RISORIA' T Has Been Thought THE HYBRID-SUBSTANCE OF THE ERYTHROCYTES OF THE HYBRIDS BETWEEN COLUMBA LIVIA AND STREPTOPELIA RISORIA’ WILMER J. MILLER2 Department of Genetics, Uni.Jersity of Wisconsin, Madison Received March 26, 1956 T has been thought that antigens of the red blood cells are more or less direct I products of their causative genes (IRWIN1939; IRWINand COLE1936a; HALDANE 1937). An exception is the “hybrid-substance” which seemingly consists of one or more antigenic specificities of the cells of hybrids between certain species of birds which are not possessed by the cells of either parental species (IRWIN1932; IRWIN and COLE1936a; IRWINand CUMLEY1945; MCGIBBON1944, 1945). The erythro- cytes of the hybrids between the common domestic pigeon, Columba lizlia, and the ring neck dove, Streptopelia risoria, possess such a hybrid-substance (IRWINand COLE193610). The segregation of this hybrid-substance in a unique backcross family and the presence or absence of related antigenic specificities in other species and certain other species hybrids of Columbidae are presented in this paper. MATERIALS AND METHODS Briefly, the procedure used to demonstrate the hybrid-substance was to produce antisera in rabbits to cells of the F1 hybrids, and then to absorb the resulting antisera at an appropriate dilution (usually 1:60) with the pooled cells of many individuals (about 10-30 each) of both parental species. If possible, the cells of the actual parents of the hybrids were used in these absorptions and as controls in the agglutination tests in order to eliminate the possibility that individual differences within either parental species might be a complicating factor in the detection of the hybrid- substance. The agglutination of the cells of the F1, and a lack of agglutination of the cells of the parental species with the resulting “reagent” were evidence for a new specificity-the “hybrid-substance”-possessed by the F1 cells. These tests were usually performed with the reagent at the absorbing dilution (1:60), and with the dilutions serially doubled, if the titer was determined. This reagent could be frac- tionated by further absorptions with cells of other species or species-hybrids which possessed serological specificities that were related to the hybrid-substance. Addi- tional details of pertinent immunological procedures such as absorption ratios of cells to serum, agglutination tests, etc., may be found elsewhere (CUMLEYand IRWIN 1941a, 1941b, 1942; IRWINand COLE1936a; MILLER1953). Most of the reagents or test fluids used in the present study were prepared from Paper No. 594 from the Department of Genetics, University of Wisconsin. This investigation was supported in part by a grant from the Research Committee of the Graduate School from funds supplied by the Wisconsin Alumni Research Foundation, and by a research grant (G-3884 to M. R. IRWIN)from the Department of Health, Education and Welfare of the Xational Institutes of Health, Public Health Service. * Present address: Serology Laboratory, School of Veterinary Medicine, University of California, Davis, California. HYBRID-SUBSTANCE IN PIGEON-DOVE HYBRIDS 70 1 four antisera (493F4, 495F6, 480F4 and 491F2) produced in rabbits against the cells of hybrids between livia and risoria. The titers of the reagents for the hybrid-sub- stance prepared from these four antisera varied from three to eight doubling dilu- tions beginning at 1: 60. Antisera against the cells of the parental species, and against cells from C/C and C/C’ backcross hybrids of C. guinea X C. livia (MILLERand BRYAN1953) were also used. Various backcross hybrids from the cross of C. guinea, the triangular spotted pigeon of Africa, with livia were available for tests. IRWIN,COLE and GORDON(1936) demonstrated the species-specific antigens A, B, C and E of guinea in contrast to livia; MILLERand BRYAN(1953) disclosed the presence of the species-specific anti- gens of livia, A’, B’, c’ and E’ which are antithetical and presumably produced by genes allelic to those producing the respective antigens of guinea. The cells of back- cross birds carrying other species-specific characters specific to S. chinensis (d-1, d-4 and d-11) and S. senegalensis (s-8 and s-10) respectively, in contrast to risoria-were included in some of these tests (see IRWIN1953 for leading references). Other char- acters specific to S. orientalis (or-4 and or-8) and S. tranquebarica (hu-4 and hu-8) have been obtained in unit form following crosses and backcrosses of each species to risoria (IRWINunpublished data). The species and F1 hybrids used were as follows: Streptopelia bitorquata (also known as dussumieri), Philippine turtle dove; S. capicola, cape turtle dove; S. chinensis, pearlneck or laceneck dove; S. orientalis, Oriental, Asiatic, or Chinese turtle dove; S. risoria, ring neck, barbary, domestic, fawn, blond or ring dove; S. semitorquata, African turtle dove; S. senegalensis, senega1 or palm turtle dove; S. tranquebarica (also known as humilis), dwarf or red backed turtle dove; S. turtur, European turtle dove; Columba jasciata, bandtail pigeon; C. Jlavirostris, yellow billed pigeon; C. guinea, triangular spotted or speckled pigeon; C. livia, common, domestic, or rock pigeon; C. maculosa, spotted pigeon, C. picazuro; C. rufina, rufous pigeon; C. palumbus, European wood pigeon; Chalcophaps indica, green wing dove; Galli- columba luzonica, bleeding heart pigeon ; Geopelia humeralis, bar-shouldered dove ; L. Cassini, cassin’s dove ; Leptotila jamaicensis, violet dove; Nesopelia galapagoensis, Galapagos dove; Ocyphaps lophotes, Australian crested dove; Phaps chalcoptera, bronze wing pigeon; Phaps elegans, brush bronze wing pigeon; Zenaida aurita, Mar- tinique dove; Zenaida asiatica, white wing dove; Zenaidura auriculata, bronze neck dove; Zenaidura graysoni, Grayson’s dove ; Zenaidura macroura, mourning dove; Caloenas nicobarica, Nicobar pigeon; Ducula spilorrhoa, white fruit pigeon; Goura cristata, Goura crowned pigeon; F1-bitorquatalrisoria; Fl-chalcoptera/elegans; FI- chinensislrisoria; Fl-chinensis/orientalis; F1-2ivia/risoria; F1-orientalislrisoria; F1- semitorquatalrisoria; F1-semitorquata/orientalis;F1-senegalensislrisoria; K-senegalen- sis/capicola; F1-tranquebarica/risoria;FI-asiatica/risoria; F1-macroura/risoria; F1- fasciata/livia; and F1-C. oenasllivia. The number of individuals used within each species varied greatly. Only one to a half dozen were available in those species diffi- cult to obtain (perhaps half of the above list). MR. J. P. TELFORDof Madison, Wisconsin, kindly allowed blood samples to be taken from his unique family of livia-risoria hybrids and backcross hybrids. MR. KARLPLATH, curator of birds, Chicago Zoological Park, kindly permitted the author 702 WILMER J. MILLER to obtain blood samples from nicobarica, spilorrhoa, lophotes and guinea; and MR. R. MARLINPERKINS, director, Lincoln Park Zoological Gardens, Chicago, Illinois, permitted the author to obtain a blood sample from cristata. RESULTS Segregation of the hybrid-substance with a species-specijic antigen Hybrids between the domestic pigeon, Columba livia, and the ring neck dove, Streptopelie risoria, are usually sterile (COLEand HOLLANDER1950). Nevertheless, MR. TELFORDraised a male F1-Zivia/risoria which in a backcross to risoria produced eight offspring which lived to maturity. One of these backcross birds produced a squab in mating to risoria. The F1 male, in addition to possessing the hybrid-sub- stance, had as expected, the species-specific characters (A’, B’, C’, and E’) of livia in contrast to guinea (MILLERand BRYAN1953). The cells of risoria lack the A’, C’ and E’ of livia, and the “B’-like” specificity which they possess was easily distin- guished from the B’ of livia (MILLER1953). Therefore, the inheritance of the hybrid- substance and the species-specific characters of livia could be followed in these eight backcross and the second backcross off spring. The data in table 1 show the segregation in the backcross offspring of the hybrid- substance and the species-specific characters of Zivia. The cells of four (Nos. 2, 6, 8 and 9) backcross birds were strongly agglutinated by any of the four reagents TABLE 1 Segregation in backcross offspring of the hybrid-substance and of characters specijc to Columba livia in contrast to Streptopeiia risoria I Presence or absence of agglutination of test cells with reagents for: Test cells (species-specific substances of lirio) hybrid-substance of Fi-liuia/risorin from antiserum B’ C’ E’ 493F4 495F6 480F4 491F2 I risoria 0 0 0 0 0 0 0 0 livia + + + + 0 0 0 0 FI-livia/rismia + + + + + + + --__+ I First backcross off- spring 1 + + 0 + 0 0 0 0 2* + + + + + + + + 3 + + 0 + 0 0 0 0 4 0 + 0 + 0 0 0 0 5 + + 0 + 0 0 0 0 6 + + + + + + + + 7 + + 0 + 0 0 0 0 8 0 0 + 0 + + + + Second backcrossoff - spring 9 + + + 0 + + + + ~- * Male parent of No. 9 in a backcross to risoria. HYBRID-SUBSTANCE IN PJGEON-DOVE HYBRIDS 703 (from antisera 493F4, 495F6, 480F4, or 491F2) for the hybrid-substance as listed in table 1. The cells of all (over 40) FI’Stested have always been strongly agglutinated by such reagents. The cells of five of the backcross offspring (Nos. 1, 3, 4, 5 and 7) and, of course, livia and risoria, including the actual risoria parent, did not react with these reagents. Each of the antigenic characters specific to livia segregated in these backcross off- spring, as expected if they were genetically determined. Thus, there were seven back- cross birds with A’ on their cells, two without; eight with and one without B’; four with and five without C’; and seven with and two without E’. There was no detect- able difference in the strength of the agglutinations of the cells of the backcross off- spring from those of the F1 parent. Of special interest is the complete correspondence in these nine birds of the presence of C’ and the hybrid-substance (Nos. 2, 6, 8 and 9 with both reactivities, and Nos. 1, 3, 4, 5 and 7 with neither). The probability that C’ and the hybrid-substance segregate independently is (99, or 1:512, if it is assumed that the genetic material responsible for the hybrid-substance does not fractionate, Or, if the 2 X 2 tabular method (FISHER 1944) is used, the probability is 1: 126.
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