Hereditas 109 151-1.57 (1988) Standard karyotype of the domestic pig Committee for the Standardized Karyotype of the Domestic Pig lngemar Gustavsson (Co-ordinator), Department of Animal Breeding and Genetics, Swedish University of Agricultural Sciences, S-7.50 07 Uppsala 7, Sweden Pig chromosomes were studied already in the beginning of thiscentury (WODSEIIAL.EK 1913). The correct chromosome number o f 38 was described as early as 1931 (KKALLIWFK 1931) but was not agreed on until the beginning of the fifties. More accurate chromosome studies co~ild.however, be carried o u t when tissue culture techniques were introduced. The first banded karyotypes occurred in 1972 (BEKGER 1072; GusrAvssoN et al. 1972; HANSEN 1972). Since then, several different banding techniques have been applied on pig chromosomes. These techniques have increased the possibilities to identify individual chromosomes as well as made possible studies of the detailed morphology. The number of known spontaneous chromosome aberrations is increasing (e.g.. POPESCII et al. 1984), and aberrations also appear to be easily induced (FKIF-s and STRANZINGER 1982). There is an extensive polymorphism (HANSEN-MELANI)LKandMti A N t l t K 1974; CHRlSTFNStNand SMEDEGARII 1978) in the heterochromatin, which appears to be of two types; G-C-rich heterochromatin, particularly located in the centromere regions of the biarmed chromosomes, and less G-Crich heterochromatin in the centromere regions o f the one-armed chromosomes (S(wNti)i.et al. 1981). There are also two nucleolnr-organizing regions, which show polymorphism ( C z A K t R and MAW 1980). The pig karyotype has recently been a popular object for gene mapping by somatic hybridization (FOKSimet a \ . 1080; <itLt.iNet a!. 1980) and the interest is growing for using in situ hybridization in the mapping work ( R A H IctN al. 1985; ECHAKI) et al. 1986). Banded karyotypes have been used for investigations of cell lines (E(w,wi) 1973) and for studies of species closely related to the domestic pig to re1078; MkI A N D t K Veal karyotype evolution ( BOSMA and HANSthi-MtI A N I ) t - K 1980). For several purposes there is thus a great need to have available a common, simple systcni f o r chromosome description and arrangement. The main identifying features of the chromosomes and an arrangcmcnt system were defined by an international study group at Rending, England, in 1976 (Proceedings of the First International Con- ference for the Standardization of Banded Karyotypes of Domestic Animals, 1980). Since then, there have been correspondence and discussions about landmark systems and more detailed chromosome descriptions at European Colloquia held in Uppsala, Sweden. in 1980. Milan-Gargnano, Italy, in 1982, and in Zurich, Switzerland, in 1984. The following description is based on those discussions. Consideration was also taken to three landmark systems and chromosome descriptions earlier proposed (HANSFN 1977; LANet al. 1980; VorcuLescuand LCNGEANLI 1980). The main objectives o f this communication have been to describe the identification banding patterns in greater detail than did the Reading report, and t o divide the chromosomes into cytologically defined regions by a landmark system which can be used for descriptions o f the normal karyotype as well as aberrant patterns. Methods The G- and R-bonding patterns of the pig chromosomes arc described by using the G T G ( S E A i m i m T 1971) and the R B A (DLlIKIi.L.AL;Xet al. 1973) banding techniques, and an R-banding technique making use o f synchronization techniques with chromosome exposure to agents which partially inhibit contraction ( R ~ N N1984). E Detailed descriptions with the band positions (but not the intensity of individual bands) are given in schematical drawings according to a computerized model kindly supplied by Dr K. Fries. Thc identification system for the bands is the same ;IS the one used for human chromosomes (Paris Conference 1972), and the landmark system also follows thc recommendations for the human nomenclature (ISCN 1978). Results Banding patterns and landmark systems agreed on arc givcn cchcmatically i n Fig. 1 and 2. Nucleolar or- Herediras 109 (1988) #I 1ti! 5 4 21 :2 21 34 43 5 6 7 3 56 12 4 #! 34 5 1 2 3 54 5 21 2 ;; 3 4 X fj! 4 7 10 1 2 43 1 12 1 2 3 4 5 1 2 423 1 2 43 423 17 3 4 5 6 65 :2 11 18 16 15 5: ?l3 9 13 Fig. 1. Diagrammatic representation of chromosome bands and landnlark system of G-banded pig chromosomes. Hereditas 10Y (1Y88J 8’-j S T A N D A R D KARYOTYPE OFTHE DOMESTIC PIG 5 4 1 2 34 2 ’1 2 56 43 457 5 4 3 l2 2’ 34 4 ;2 f l 5 2 34 5 54 3 Bi 3 4 5 1 0 47 ill! 4 125 17 18 n F 13 Fig. 2. Diagrammatic reprewntation o f chromosome hands and landmark \?stem of R-banded pig chromosomes. 153 154 I . GUSTAVSSON Fig. 3. Representative GTG-banded male pig karyotype (courtesy I . Gustavsson). Hereditas I09 (1988) Hereditas 109 (1988) STANDARD KARYOTYPE OF THE DOMESTICPIG 155 Fig. 4. Representaive R-banded female pig karyotype after syncronization with rnethotrexate-leucovorin and fluorouracil. release of the synchronization block with 5-brornodeoxyuridine and partial inhibition of contraction with Hoechst 33258 (courtesy M. R ~ n n e ) . ganizer regions (NORs) occur in the centromeric parts of the p arms of chromosomes 8 and 10. Representative karyotypes are demonstrated in Fig. 3 and 4 and the bands serving as landmarks are described in Table 1 . Conclusion Although there is today an increased usage of highresolution techniques, it was considered necessary first to describe those banding patterns which can be 156 Heredirus I09 (1988) I GUSTAVSSON l i r h l i ~ 1 . Gliiinck \civing ;I\ landmiiih\ in ihe pig \tendard kiiryiitype (iinii\\iiin iif ii chromo\omc o r ii chromosome arm indicates that either both a r m \ or the arm in queatiim consi5th iif only o n e region) Chromowmc no. I Aim Numhcrof Landmarks region\ 2 3 Central negative band (21) Central negative band (21) Broad proximal negative hand (71) Central negative hand (21) Negative b a n d ( 2 l ) i n the distal half Central negative band (21) Proximal broad negative band (21), negative band in the distal half (31) 7 Central strong positive band (21) X Centralnegative band(21) Broad negative band in proximal half (2 I ) 9 Positive band in distal half (21) Centralpositive band (21) 13 Proximal negative (21) and cential negativeband (31). negative band in the diatal half (41) I4 Negative hand in the pioximal haif ( 2 I ) IS Central positive band (21) 16 Broad negative band in the dim1 half(21) 17 Broad di\tal negative hand ( 2 I ) 1x Po\itivecentral band (21) X Positive central band (21) Proximal positive band (21) identified in moderately contracted chromosomes. The banding patterns of the chromosomes and the landmark systems presented in this work make possible accurate description of small chromosome segments. This work should, however, be continued with the establishment of a high resolution karyotype. Literature cited BERGER. K. 1972. h d e du caryotype du porc avec une nouvelle technique. - Exp. Cell He$. 75: 29X-3IHJ BOSMA, A . A . 1Y7R. T h e c h r o m o w n a l G h i n d i n g pattern in the ~. and its wart hog. Phucochoerirs wrhiopicus ( S u i d ; ~M;immalia) implication$ f o r the sy\tematic po\ition 01 the \pecies Geneticu 49: 1S- I9 CHRISTENSEN, K. and SMEDECAKI). K 1Y7X Chromosome marker in domestic pigs. C-band polymorphivn -Herediru,s 88: 269-272 ('/.AKt-K. R. and MAYK. B . I W I . Derccrznn o f nuclcolar orgiinizer region\ in the chrnmosomea ol t h e domestic pig (Swi ,si.ro/u donw.sriu~L . ) - Experirnrru 30: 135&1357 DUTKILLAUX.R.. LAUREN'I, C.'.. COUTIJRIEK, J . a n d L E J E U N E ,J . 1Y73. Coloration d e s chromosomes humains p a r I'acridine orange iiprcs traitement p a r le 5-bromodeoxyuridine. -C. R. Acod .Sci. (1'uri.s) 270: 3179-31x1 ECHAKD.G . 197.1. f t u d e d e \ h m d e s chromosomiques du porc et d e trois diffcrente\ s o w h e \ d e rein d e porc e n culture ( P K I S . F Pf KP). -Ann. (;Cnt'f Sd. .4fllm. S: 1-21 EC'HAKD.G..YtKl.t..J..(jtl I I N . J .DAl.ENS.M.andGILl.Olb. M . 19x6. Asvgnment of the major histocompatibility complex t o the pl .&ql.2 region o f chromosome 7 in the pig (Sus scrofu r/onie,sricu 1- ) hv i n \itu hyhridization. - Cyrogener. Cel/ Gener. 41: 126-12x F 0 R S l ' t . H . M . . SIKAN/.IN(;€K. G dnd H E L L K U H L B . IYXO Xchromoscimc gene awignment 01 \wine a n d cattle. - Nanrrwissrnschufirn 67:4 x 4 9 FRIES, K.and s'rKAN7lNGEK. G . 1982. Chromosomal mutations Cyrogenet. Cell in pigs derived from X-irrkidiated semen. Genrr. 34: S 5 4 t i GELL.IN. J . , BENNE, F . HOKS-CAYLA. M . C. and GILLOIS. M . IYXO. C a r t c gknique du porc ( S u s w o f u L . ) . I . etude d e deux groupe\ \ y n t h i q u e (ihPI). PGK. H P R T e l PMK2. M P I . Ann. GPnCf 23: 15-21 (;I'SIAVSSON. I . . H A M I I O K N . kl . J O H A N S S O N , C a n d Z t ( ' H . L 1Y72. Identilic;iti(in ot the pig chroinosomcr by the quinacrine ~ H(wdrto\ 100 ( I Y X X j STANDARD KAKYOTYPE OFTHE DOMESTICPIG niu.rtard fluorc\cence technique. - E x / ) . Cell Kes. 70: 471-474 HANSEN. K . M. 1972. The karyotype 01 the pig(Sus scrofl] dome.crim).identified by qiiinxrinc mu\t;ird staining and fluorescence ( ~ y r ~ i ~ i wI rI .~ 2x6 i ~ \ ?Y4 micro\copy. HANSEN. K . M . 1977. Identification o f the chromostime\ of the domestic pig (Sir5 ,\crofu dorric~.\tfca).An identification key and a kindmark system - A n i i GPnc'r. Srl. Anim. Y: 517-ti26 HANSEN-MELANDt.K. E. ;ind MILANDER, Y . 1974. The ~ I4')-15X karyotype 0 1 thc pig. - t l e r c ~ i l i f u77 ISCN IY7X. A n international rystcm for human cytogenetic nomenclature. - ('yrogrrier ( ' d l Genet. 21: 309-404 KRAI.I.INGER. H. F. 1931. Cytologische Studien in einigen Haus\;iugctieren. H~ihilitationssctirift. Arch. f. Tirrrmuhrcing irnrl / ' ~ w z i i i ~ / Ah!. it. H5 Brc,\liiir L l N . C. C.. BIEDERMAN. B. M.. JAMRO, H . K . . HAWTHORNE. A . B. and C H U R C H .R. B. 1980. Porcine (Sus tcrofu domesrico) chromosome identification and \uggcsted nomenclature. cuff..I. Grncr. < j t o l . 22. 103-1 I 6 Mt.I.Ahl)LK. Y . and HANS1 N-Mt.1 A h l X K , E . 19XO. C'hromohome 5tudies in African wild pig\ (Suidae, Mammalia). Heredituc Y2: 283-280 Paris Conference 1Y72. Standardization i n human cytogenctics. Cytogmeric~I I : 3 13-362 POI'CSC'U. C'. P.. BONNEAI , M , 'I I X I L K . M . . BAHKI. I . and BOSCHER. J . 1984. Kecipmcal tran4ocations in pigs. Their detection and consequence>o n animal performance and economic Iosse$. -J. Hcrcrl. 75: 4 4 W 5 2 Proceeding of the Fint International Conference for the Standardi u t i o n ol Banded K;iry<rtype\ 0 1 I)wncatic Animal\ Reading, England. 1976. (Eds. C. E . FORD. D . L. POLI.OCK und 1. GUSTAVSSO3). 1980. - Hcredrms 92: 145-162 KAIHN. M . . FKIES. R . , SINGI:K, D. and K1JDI)I.E. F. H . 1Y8.5. Aa\ignmcnt of the porcine major histocompatibility complex t o . Giwt,t. chr~imos(ime7 hy in a i t u hyhridizatim - C ' y r o ~ e w r (LII 39: 20&20Y KDNNE. M. l Y X 4 . Fluorouracil synchronization of pig lymphocyte cultures. Induction of high resolution K-handing by in vitro exposure to S-bromodeoxyuridine/Hoech\t33258. -6th Eur. Colb ~ q (:~rogc,~fc't . L)<IWW.S/A I I I I I I .I Y W , p. 1%-196 <CHNEDL. W . . ABKAHAM, R . , FOIISTEK. M . and SCHWEIZER. D . IYXl. Differential fluorevxiit staining of porcinc heterochromatin by chrcimomycin A3/di\tamycin AiDAPl and D287/ 170. - Cyrogenet. C d l ( h e r 3 / :249-253 ~ ~ 157 M . 1Y71 A rapid banding technique tor human chriiiii~i\onie\.- Liiricer (2): 971-972 VOII'LiI.ESC"J. 1. and LUNGEANU.A . 1980. Some comments o n the G-hand\ i n pig chromosome\. -4th Eicr. Colloq. Cyrogenet. I h m i w Aninr. IYXfI, p. 363-370 WODSEDALEK.J . E . 1913. Spermatogenesk of the pig with specia1 reference to the accessory chromosomes. - B i d . Bull. 25: 8-32 StAHKIC;III. Members of the committee B A H R I .I . Lahoratoirc de Cytogenttique. UNCEIA-INRA, Recherches Zi)otechniquea. 78350 Jouy-en-Josas, France BOSMA. A . A . Drp;irrment of Functional Morphology, Faculty of Veterinary Science\. State Univervty. Yalelaan 1,3508 TD, Utrecht. The Netherl;rnds ECHARD,G . Lahoratoire de Gknetique Cellulaire. Centre de Recherchea de Toulou\e, INKA, BP 27, 31326 Castanet-Tolosan. France FRIF,S,R. lnstitut fur Tierproduktion, Gruppe Tierzucht, ETHZent rum, XOY? Zurich, Switzci-land GUSI'AVSSON.I . (Co-ordinator). Department of Animal Breeding and Genetic\. Swedish University of Agricultural Sciences, S-7.50(17 Uppsala 7. Sweden HAAN. N. A . I)E. Department of Functional Morphology, Faculty <it Veterinary Science\, State Univerrity. Yalelaan I . 3508 'ID. Utrecht. The Nctherlands HANSEN, E. M . University Mcd. Anatomy. Dept. B, Panum Inst., Blegdamsvcj 3. DK-ZZOO Copenhagen, Denmark HANSEN.K M. University Med. Anatomy, Dept. B. Panum Inat., Blogdani\vej 3. DK-22OO ('openhagcn. Denmark M A K I N E NA . Department of Animal Breeding and Genetics, Swedish Univer5ity of Agricultural Sciences, S-75007 Uppsala 7, Sweden POPESCU. C. P. Laboratoirc de Cytogenetique, UNCEIA-INRA. Centrc National de Recherche3 Zootechnique\ 7x350 Jouy-enJojas, France KDNNE, M . Univenity of Odense. Winslow Institute of Human Anatomy. DK-S230 Odense, Denmark SYSA. P. Department of Histology and Embryology, Warsaw Agricultural Univervty. 2-766 Warsaw. Poland TROSHINA, A . Imtitute of Cytology and Genetics. Novosibirsk63owo. USSK
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