AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 85:335-338 (1991)
Brief Communication: Immunoglobulin (Gm and Km) Allotypes in Two Populations of Catalonia (Spain) PASCUAL MORENO AND HIDE0 MATSUMOTO Department of Animal Biolo ylSection of Anthropology, Facult of Biology, University of Barcefne, 08028 Barcelona, Spain ( P . 4 ; Department of Legal Medicine, Osaka Medical School, Duigukumachi, Takatsuki, Osaka 569, Japan (H.M.)
KEY WORDS
Population genetics, Immunoglobulin allotype
ABSTRACT
Serum samples from two populations of Catalonia, Spain, 208 from Olot (Gerona) and 209 from Tortosa (Tarragona), were typed for Glm (1, 2,3,17),G3m (5,10,11,13,14,15,16,26),and Km (1).The Gm patterns ofthe Catalonian populations are characterized by the presence of four haplotypes, Gm 1,17;21,26 Gm 1,2,17;21,26 Gm 1,3;5,10,11,13,14,26and Gm 3;5,10,11, 13,14,26. The homogeneity for haplotype Gm 1,17;21,26among our data and other European populations suggests the existence of an isofrequency line which starts from the Mediterranean zone of Iberian Peninsula and continues through the northwestern part of Europe. From this line a decreasing cline towards the south can be observed. For the haplotype Gm 1,2,17;21,26, affinities are observed between Catalonian populations and other populations from central Europe. This confirms the existence of a gradient towards low values from NW to SE. The presence of the typical Mongoloid haplotype Gm 1,3;5,10,11,13,14,26is discussed in this paper. No significant differences in the frequencies of the Kml allele were observed among the European populations.
The extensive range of allotypic antigens to the De artment of Legal Medicine, Osaka, present in human immunoglobulins furnish Japan, w ere the Gm and Km typings were useful evidence on genetic variability among carried out. All the samples were typed for and within human populations. Three major Glm (1,2, 3, 17), G3m (5,10, 11, 13, 14, 15, systems are currently recognized: Gm anti- 16,21,26) and Km (1).The reagents used are gens are characteristic of y heav chains, Am presented in Table 1.Haplot e fre uencies anti ens of a heavy chains, and k n antigens and degree of fit with the grdy-deinberg of K ht chains. equilibrium were determined (in Osaka) usAltaough the distribution of human im- ing the com uter program MAXIM (Kurcmunoglobulin allotypes has been analyzed in zynski and teinberg, 1967). several European populations, little is RESULTS AND DISCUSSION known in this regard about the Spanish Gm system opulation. In this a er we present the The phenotype and the haplotype fre uenEndings on Gm and &laplotypes of serum sam les from two populations of Catalonia cies in the two populations analyze! are ( N q Spain). The two populations are from shown in Table 2 and Table 3. Six different Olot in the north ( rovince of Gerona) and Gm phenot pes were found, which can be Tortosa (province o Tarragona) in the south. explained y four haplotypes: Gm 1,17; 21,26, Gm 1,2,17;21,26, Gm 1,3;5,10,11,13, MATERIAL AND METHODS 14,26, and Gm 3;5,10,11,13,14,26. The disSerum samples from a total of 417 unre- tribution of phenotypes is in good agreement lated and healthy individuals (both sexes) from two PO ulations of Catalonia, 208 from Olot and 20 from Tortosa, were collected in Received October 4, 1989; accepted January 16, 1991 early 1988. These samples were air-mailed
K
rp
P
8
@
1991 WILEY-LISS. INC.
i
336
P. MORENO AND H. MATSUMOTO
with the Hardy-Weinber law, as shown by the test for goodness o fit (Table 2). T i e comparison (by means of the x2 test) of the two Catalonian populations, taking into consideration the four Gm haplotypes already analyzed, shows a clear homo eneity. When comparisons are carried out or each haplotype se arately, the homogeneit persists for all o them except for the hap otype Gm 1,3;5,10,11,13,14,26,which reveals significant differences between the two populations (x2(1,= 4.61; P = 0.032). Our attention is drawn to the presence of the haplotype Gm 1,3;5,10,11,13,14,26in the
f
f
P
i?
TABLE 1. Reagents used for G m and K m allotype determinations
Aluhabetic
Numerical
Antiallotwe
Anti-Rho
Glm a
1
X
2 3 17
3552 2984 2871 3272
2880 2880
5 11 13 14 10 21 15 16 26
7514 0058 4721 0663 1340 1642 2624 5198 1369
3656 3656 3656 3656 3656 3359 3068 3068 Eggen
1
5872
2447
f 2
G3m bl b0 b3 b4 b5 g S
t U
~~~~
~~
~
KO-Ro 2880
Km I
~
two Catalan populations studied. Frequencies of this are highest in the Mongoloid eoples of East Asia and lowest in the west of Europe. Walter et al. (1987) analyzed ten endogamous PO ulations of Assam, India (Caucasoid a n 8 Mongoloid r p s ) , and found that the frequencies o f t is haplotype vary from 0.127 to 0.449 in the Caucasoid On the other hand, Hollan and ublished data) have observed a 0.0308 in Hungarians, and frequencies of 0.1514 and 0.2133 in two groups ofHungarian Gypsies.The two Catalan populations show very low frequencies of this haplotype (0.0461 and 0.0196). These data sugest the existence of a long-standing enetic ow from East Asia to Europe. More ata on other European po ulations are needed to confirm this possib e explanation. The comparison of our results with those of other Euro ean populations can take into account ony the ha lotypic frequencies of Gm 1,17;21,26 and 8 m 1,2,17;21,26, since not all opulations have been tested for the same a otypes. In order to simplify the comparisons, we have combined the two Catalonian populations into a single group since there are no significant differences between them for the two haplotypes cited above. In consequence the Catalonian population is represented by these frequencies: Gm 1,17;21,26 = 0.2415 and Gm 1,2,17;21,26 = 0.0463. With regard to haplotype Gm 1,17;21,26, the frequency for the Catalonian population
8
%
P
P
R
TABLE 2. G m phenotypes in Catalonian populations: conformity with Hardy- Weinberg expectations
Olot (Gerona)’ Observed Expected
Phenotype 1,17;21,26 1,2,17;21,26 1,3,17;5,10,11,13,14,21,26 1,2,3,17;5,10,11,13,14,21,26 1,3;5,10,11,13,14,26 3;5,10,11,13,14,26 Tohl iX,” = 4.576 df X = 2.535; df
= =
14 1
72 17 13 91 208
12.2 4.8 72.0 12.9 13.3 92.8 208.0
Tortosa (Tarragona)’ Observed Expected 16 6 63 14 6 104 209
12.1 5.4 71.4 14.6 5.8 99.8 209.0
2; probability = 0.102. 2; probability = 0.282.
TABLE 3. G m halotype frequencies in Catalonian populations
Haplotype 1,17;21,26 1,2,17;21,26 1,3;5,10,11,13,14,26 3;5,10,11,13,14,26
Olot (Gerona) Frequency 0.2426 0.0435 0.0461 0.6678
SE 0.0211 0.0101 0.0123 0.0241
Tortosa (Tarragona) Frequence SE 0.2403 0.0492 0.0196 0.6909
0.0210 0.0107 0.0081 0.0230
337
IMMUNOGLOBULIN ALLOTYPES IN CATALONlA
does not differ significantly from the population of Valencia (x2(1,= 0,796; P = 0.3723) (Shanfield et al., 1981) nor from such other north and west Euro ean populations as Scotland (Francis et ,a! 1986), Great Britain, Norway, Finland, and France (Johnson et al., 1977a).On the other hand, the Catalonian data exhibit marked and statistically significant differences from data from Portugal (Pandey et al., 19821, several Italian series (Steinberg et al., 19811,two Romanian series (Johnson et al., 1977b) one Dutch series (Biemond et al., 1987), and several series from central Europe representing populations of Czechoslovakia, Austria, Germany, Poland and Hungary (Schanfield et al., 1975). The homogeneity of the frequency of haplotype Gm 1,17;21,26 among the two series sampled for this study and the series of Valencia evidences a uniformity within the Mediterranean zone of the Iberian Peninsula. The marked differences amon the above mentioned groups and Portuga suggest that there is a clear differentiation between the Mediterranean and Atlantic zone of the Iberian Peninsula. In studies of some other polymorphisms-for instance, GC and ESD-significant differences between Catalonian populations and Portugal have also been observed (Moral, 1986). The coincidence of our results with the data of other northern and southern Euroean populations (Finland, Norway, Scotrand, Great Britain, and France) suggests the existence of an isofrequency line which starts from the Mediterranean zone of the Iberian Peninsula and continues across the west and north part ofEurope. From this line a decreasing cline towards the south can be observed, as pointed out b Grubb (1961). Comparison of the Cata onian series with the series cited above for ha lotype Gm 1,2,17;21,26 reveals results di ferent from those obtained for the haplotype Gm 1,17;21,26: similarities with central European populations (Hungarian, Romanian, Polish, and Czechoslovak series) are observed. For the haplotype Gm 1,2,17;21,26, we thus find a European gradient from high values in the northwest toward low values in the southeast, as suggested by Steinberg (1981). Km system Km phenotypes and allele frequencies of the two Catalonian populations are given in Table 4.No significant differences have been
4
Y
P
TABLE 4. K m phenotype and allele frequencies among two poDulations
Olot (Gerona) Km phenotype 1+ 1Total Km allele frequency Kml
Tortosa (Tarragona)
35 173 208
31 179 210
0.088
0.077
found from other eninsular populations such as Barcelona Hernandez, 19811, Valencia (Schanfield et al., 1981) ar Madrid (Llorente et al., 1976).The comparison of our samples with other European populations like Austria, Czechoslovakia, Finland, Germany, Greece, Hungary, Ireland, Italy, Norway, Poland, Switzerland, and Yugoslavia (Steinberg, 1981) demonstrates a homogeneity for this system throug out Europe. The Km locus is thus not especially useful for discriminating among European populations.
P
rat
ACKNOWLEDGMENTS
We ex ress our gratitude to Dr. Miquel Ris au ( 0s ital de Sant Jaume de Olot) and to r. Joan aragoza (Hospital Ntra. Sra., de la Cinta de Tortosa) for his invaluable help in collecting the serum samples.
8
LITERATURE CITED Biemond I, Lange G, Weterman I, and Pena AS (1987) Immunoglobulin allotypes in Crohn's disease in the Netherlands. Gut. 28:610-612. Francis DA, Brazier DM, Batchelor JR, McDonald WI, Downie AW, and Hern JEC (1986) Gm allotypes in multiple sclerosis: Influence susceptibility in HLADQwl-positive patients from the north-east of Scotland. Clin. Immunol. Immunopathol. 41:409-416. Grubb R (1961) Hereditary gamma globulin groups in man. Am. J. Hum. Genet. 13:171-174. Hernandez M (1981)El factor serico Inv(1)en Barceloneses. Rev. Mex. Estud. Antrop. xxvll:1:81-88. Johnson WE, Kohn PH, and Steinberg AG (1977a) Pooulation eenetics of the human allotvDes Gm. Inv and h m .An-analytical review. Clin. Immunol. immunopathol. 7:97-113. Johnson WE. Kohn PH. and Steinbere AG (197713)Gm and Km(Inv) frequencies in two Roumanian populations. Hum. Genet. 39:199-211. Kurczynski TW, and Steinberg AG (1967) A general r g r a m for maximum likelihood estimation of gene requencies. Am. J. Hum. Genet. 19:178-179. Llorente L, Campillo FL, Gallardo LE, Cuesta JA, and Senra A (1976)Distribucion de 10s grupos Gml, Gm2, GmlO e Invl, en la poblacion espanola. Sangre 21(11:87-89.
338
P. MORENO AND H. MATSUMOTO
Moral P (1986)Estudio antropogenetico de diversos polimorfismos hematologicos en la isla de Menorca. D. Thesis, University of Barcelona. Pandey JP, Shannon BT, Arala-Chaves MP, and Fudenberg HH (1982)Gm and Km frequencies in a Portugese population. Hum. Genet. 61:154-156. Shanfield MS, Herzog P, and Fudenberg HH (1975) Immunoglobulin allotypes of European Populations. 11.Gm, Am and Krn(1nv)allotypic markers in Czechoslovakians. Hum. Hered. 25382-392. Schanfield MS, Baylerian R, Maiquez J, and Carbonell F
(1981) Immunoglobulin allot pes in European populations. IV. Gm, Am and Km alfotypic markers in Valencia, Spain. J. Immunogenet. 8.529-532. Steinberg AG, and Cook CE (1981) The distribution of the Human Immunoglobulin Allotypes. Oxford University Press. Walter H, Matsumoto H, Mi azaki T, Mukherjee BN, Malhotra KC, Das BM, Gilgert K, and Lindenberg P (1987)Distribution ofGm andKm allotypes among ten populations of Assam, India. Am. J. Phys. Anthropol. 73:439-445.
Brief Communication: Immunoglobulin (Gm and Km) Allotypes in Two Populations of Catalonia (Spain) PASCUAL MORENO AND HIDE0 MATSUMOTO Department of Animal Biolo ylSection of Anthropology, Facult of Biology, University of Barcefne, 08028 Barcelona, Spain ( P . 4 ; Department of Legal Medicine, Osaka Medical School, Duigukumachi, Takatsuki, Osaka 569, Japan (H.M.)
KEY WORDS
Population genetics, Immunoglobulin allotype
ABSTRACT
Serum samples from two populations of Catalonia, Spain, 208 from Olot (Gerona) and 209 from Tortosa (Tarragona), were typed for Glm (1, 2,3,17),G3m (5,10,11,13,14,15,16,26),and Km (1).The Gm patterns ofthe Catalonian populations are characterized by the presence of four haplotypes, Gm 1,17;21,26 Gm 1,2,17;21,26 Gm 1,3;5,10,11,13,14,26and Gm 3;5,10,11, 13,14,26. The homogeneity for haplotype Gm 1,17;21,26among our data and other European populations suggests the existence of an isofrequency line which starts from the Mediterranean zone of Iberian Peninsula and continues through the northwestern part of Europe. From this line a decreasing cline towards the south can be observed. For the haplotype Gm 1,2,17;21,26, affinities are observed between Catalonian populations and other populations from central Europe. This confirms the existence of a gradient towards low values from NW to SE. The presence of the typical Mongoloid haplotype Gm 1,3;5,10,11,13,14,26is discussed in this paper. No significant differences in the frequencies of the Kml allele were observed among the European populations.
The extensive range of allotypic antigens to the De artment of Legal Medicine, Osaka, present in human immunoglobulins furnish Japan, w ere the Gm and Km typings were useful evidence on genetic variability among carried out. All the samples were typed for and within human populations. Three major Glm (1,2, 3, 17), G3m (5,10, 11, 13, 14, 15, systems are currently recognized: Gm anti- 16,21,26) and Km (1).The reagents used are gens are characteristic of y heav chains, Am presented in Table 1.Haplot e fre uencies anti ens of a heavy chains, and k n antigens and degree of fit with the grdy-deinberg of K ht chains. equilibrium were determined (in Osaka) usAltaough the distribution of human im- ing the com uter program MAXIM (Kurcmunoglobulin allotypes has been analyzed in zynski and teinberg, 1967). several European populations, little is RESULTS AND DISCUSSION known in this regard about the Spanish Gm system opulation. In this a er we present the The phenotype and the haplotype fre uenEndings on Gm and &laplotypes of serum sam les from two populations of Catalonia cies in the two populations analyze! are ( N q Spain). The two populations are from shown in Table 2 and Table 3. Six different Olot in the north ( rovince of Gerona) and Gm phenot pes were found, which can be Tortosa (province o Tarragona) in the south. explained y four haplotypes: Gm 1,17; 21,26, Gm 1,2,17;21,26, Gm 1,3;5,10,11,13, MATERIAL AND METHODS 14,26, and Gm 3;5,10,11,13,14,26. The disSerum samples from a total of 417 unre- tribution of phenotypes is in good agreement lated and healthy individuals (both sexes) from two PO ulations of Catalonia, 208 from Olot and 20 from Tortosa, were collected in Received October 4, 1989; accepted January 16, 1991 early 1988. These samples were air-mailed
K
rp
P
8
@
1991 WILEY-LISS. INC.
i
336
P. MORENO AND H. MATSUMOTO
with the Hardy-Weinber law, as shown by the test for goodness o fit (Table 2). T i e comparison (by means of the x2 test) of the two Catalonian populations, taking into consideration the four Gm haplotypes already analyzed, shows a clear homo eneity. When comparisons are carried out or each haplotype se arately, the homogeneit persists for all o them except for the hap otype Gm 1,3;5,10,11,13,14,26,which reveals significant differences between the two populations (x2(1,= 4.61; P = 0.032). Our attention is drawn to the presence of the haplotype Gm 1,3;5,10,11,13,14,26in the
f
f
P
i?
TABLE 1. Reagents used for G m and K m allotype determinations
Aluhabetic
Numerical
Antiallotwe
Anti-Rho
Glm a
1
X
2 3 17
3552 2984 2871 3272
2880 2880
5 11 13 14 10 21 15 16 26
7514 0058 4721 0663 1340 1642 2624 5198 1369
3656 3656 3656 3656 3656 3359 3068 3068 Eggen
1
5872
2447
f 2
G3m bl b0 b3 b4 b5 g S
t U
~~~~
~~
~
KO-Ro 2880
Km I
~
two Catalan populations studied. Frequencies of this are highest in the Mongoloid eoples of East Asia and lowest in the west of Europe. Walter et al. (1987) analyzed ten endogamous PO ulations of Assam, India (Caucasoid a n 8 Mongoloid r p s ) , and found that the frequencies o f t is haplotype vary from 0.127 to 0.449 in the Caucasoid On the other hand, Hollan and ublished data) have observed a 0.0308 in Hungarians, and frequencies of 0.1514 and 0.2133 in two groups ofHungarian Gypsies.The two Catalan populations show very low frequencies of this haplotype (0.0461 and 0.0196). These data sugest the existence of a long-standing enetic ow from East Asia to Europe. More ata on other European po ulations are needed to confirm this possib e explanation. The comparison of our results with those of other Euro ean populations can take into account ony the ha lotypic frequencies of Gm 1,17;21,26 and 8 m 1,2,17;21,26, since not all opulations have been tested for the same a otypes. In order to simplify the comparisons, we have combined the two Catalonian populations into a single group since there are no significant differences between them for the two haplotypes cited above. In consequence the Catalonian population is represented by these frequencies: Gm 1,17;21,26 = 0.2415 and Gm 1,2,17;21,26 = 0.0463. With regard to haplotype Gm 1,17;21,26, the frequency for the Catalonian population
8
%
P
P
R
TABLE 2. G m phenotypes in Catalonian populations: conformity with Hardy- Weinberg expectations
Olot (Gerona)’ Observed Expected
Phenotype 1,17;21,26 1,2,17;21,26 1,3,17;5,10,11,13,14,21,26 1,2,3,17;5,10,11,13,14,21,26 1,3;5,10,11,13,14,26 3;5,10,11,13,14,26 Tohl iX,” = 4.576 df X = 2.535; df
= =
14 1
72 17 13 91 208
12.2 4.8 72.0 12.9 13.3 92.8 208.0
Tortosa (Tarragona)’ Observed Expected 16 6 63 14 6 104 209
12.1 5.4 71.4 14.6 5.8 99.8 209.0
2; probability = 0.102. 2; probability = 0.282.
TABLE 3. G m halotype frequencies in Catalonian populations
Haplotype 1,17;21,26 1,2,17;21,26 1,3;5,10,11,13,14,26 3;5,10,11,13,14,26
Olot (Gerona) Frequency 0.2426 0.0435 0.0461 0.6678
SE 0.0211 0.0101 0.0123 0.0241
Tortosa (Tarragona) Frequence SE 0.2403 0.0492 0.0196 0.6909
0.0210 0.0107 0.0081 0.0230
337
IMMUNOGLOBULIN ALLOTYPES IN CATALONlA
does not differ significantly from the population of Valencia (x2(1,= 0,796; P = 0.3723) (Shanfield et al., 1981) nor from such other north and west Euro ean populations as Scotland (Francis et ,a! 1986), Great Britain, Norway, Finland, and France (Johnson et al., 1977a).On the other hand, the Catalonian data exhibit marked and statistically significant differences from data from Portugal (Pandey et al., 19821, several Italian series (Steinberg et al., 19811,two Romanian series (Johnson et al., 1977b) one Dutch series (Biemond et al., 1987), and several series from central Europe representing populations of Czechoslovakia, Austria, Germany, Poland and Hungary (Schanfield et al., 1975). The homogeneity of the frequency of haplotype Gm 1,17;21,26 among the two series sampled for this study and the series of Valencia evidences a uniformity within the Mediterranean zone of the Iberian Peninsula. The marked differences amon the above mentioned groups and Portuga suggest that there is a clear differentiation between the Mediterranean and Atlantic zone of the Iberian Peninsula. In studies of some other polymorphisms-for instance, GC and ESD-significant differences between Catalonian populations and Portugal have also been observed (Moral, 1986). The coincidence of our results with the data of other northern and southern Euroean populations (Finland, Norway, Scotrand, Great Britain, and France) suggests the existence of an isofrequency line which starts from the Mediterranean zone of the Iberian Peninsula and continues across the west and north part ofEurope. From this line a decreasing cline towards the south can be observed, as pointed out b Grubb (1961). Comparison of the Cata onian series with the series cited above for ha lotype Gm 1,2,17;21,26 reveals results di ferent from those obtained for the haplotype Gm 1,17;21,26: similarities with central European populations (Hungarian, Romanian, Polish, and Czechoslovak series) are observed. For the haplotype Gm 1,2,17;21,26, we thus find a European gradient from high values in the northwest toward low values in the southeast, as suggested by Steinberg (1981). Km system Km phenotypes and allele frequencies of the two Catalonian populations are given in Table 4.No significant differences have been
4
Y
P
TABLE 4. K m phenotype and allele frequencies among two poDulations
Olot (Gerona) Km phenotype 1+ 1Total Km allele frequency Kml
Tortosa (Tarragona)
35 173 208
31 179 210
0.088
0.077
found from other eninsular populations such as Barcelona Hernandez, 19811, Valencia (Schanfield et al., 1981) ar Madrid (Llorente et al., 1976).The comparison of our samples with other European populations like Austria, Czechoslovakia, Finland, Germany, Greece, Hungary, Ireland, Italy, Norway, Poland, Switzerland, and Yugoslavia (Steinberg, 1981) demonstrates a homogeneity for this system throug out Europe. The Km locus is thus not especially useful for discriminating among European populations.
P
rat
ACKNOWLEDGMENTS
We ex ress our gratitude to Dr. Miquel Ris au ( 0s ital de Sant Jaume de Olot) and to r. Joan aragoza (Hospital Ntra. Sra., de la Cinta de Tortosa) for his invaluable help in collecting the serum samples.
8
LITERATURE CITED Biemond I, Lange G, Weterman I, and Pena AS (1987) Immunoglobulin allotypes in Crohn's disease in the Netherlands. Gut. 28:610-612. Francis DA, Brazier DM, Batchelor JR, McDonald WI, Downie AW, and Hern JEC (1986) Gm allotypes in multiple sclerosis: Influence susceptibility in HLADQwl-positive patients from the north-east of Scotland. Clin. Immunol. Immunopathol. 41:409-416. Grubb R (1961) Hereditary gamma globulin groups in man. Am. J. Hum. Genet. 13:171-174. Hernandez M (1981)El factor serico Inv(1)en Barceloneses. Rev. Mex. Estud. Antrop. xxvll:1:81-88. Johnson WE, Kohn PH, and Steinberg AG (1977a) Pooulation eenetics of the human allotvDes Gm. Inv and h m .An-analytical review. Clin. Immunol. immunopathol. 7:97-113. Johnson WE. Kohn PH. and Steinbere AG (197713)Gm and Km(Inv) frequencies in two Roumanian populations. Hum. Genet. 39:199-211. Kurczynski TW, and Steinberg AG (1967) A general r g r a m for maximum likelihood estimation of gene requencies. Am. J. Hum. Genet. 19:178-179. Llorente L, Campillo FL, Gallardo LE, Cuesta JA, and Senra A (1976)Distribucion de 10s grupos Gml, Gm2, GmlO e Invl, en la poblacion espanola. Sangre 21(11:87-89.
338
P. MORENO AND H. MATSUMOTO
Moral P (1986)Estudio antropogenetico de diversos polimorfismos hematologicos en la isla de Menorca. D. Thesis, University of Barcelona. Pandey JP, Shannon BT, Arala-Chaves MP, and Fudenberg HH (1982)Gm and Km frequencies in a Portugese population. Hum. Genet. 61:154-156. Shanfield MS, Herzog P, and Fudenberg HH (1975) Immunoglobulin allotypes of European Populations. 11.Gm, Am and Krn(1nv)allotypic markers in Czechoslovakians. Hum. Hered. 25382-392. Schanfield MS, Baylerian R, Maiquez J, and Carbonell F
(1981) Immunoglobulin allot pes in European populations. IV. Gm, Am and Km alfotypic markers in Valencia, Spain. J. Immunogenet. 8.529-532. Steinberg AG, and Cook CE (1981) The distribution of the Human Immunoglobulin Allotypes. Oxford University Press. Walter H, Matsumoto H, Mi azaki T, Mukherjee BN, Malhotra KC, Das BM, Gilgert K, and Lindenberg P (1987)Distribution ofGm andKm allotypes among ten populations of Assam, India. Am. J. Phys. Anthropol. 73:439-445.
