Standard karyotype of the domestic pig

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.
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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