Molecular analysis of esterase D polymorphism

We have analyzed the esterase D (EsD) polymorphism at the nucleic acid level. Two common alleles, EsD¹ and EsD², are characterized by the substitution of one amino acid (Gly-to-Glu), which is caused by the point mutation of one nucleotide (G-to-A). Individuals exhibiting the EsD1 and EsD 2 phenotypes are homozygotes for EsD 1 and EsD 2 cDNAs, respectively. Individuals showing the EsD 2-1 phenotype have two kinds of cDNAs, viz., EsD 1 and EsD 2. The point mutation difference between the cDNAs of the EsD¹ and EsD² alleles results in a different SspI digestion site. A restriction fragment length polymorphism caused by this difference with respect to the SspI digestion site makes it possible to determine the EsD phenotype using DNA samples extracted from forensic materials with no EsD enzymatic activity.     

Isoelectric focusing studies of human red cell esterase D: Evidence for polymorphic occurrence of a new allele EsD 7 in Japanese

Summary The isoelectric focusing study of esterase D in Japanese revealed evidence of a new polymorphic allele (EsD ⁷) which is difficult to find by conventional starch gel electrophoresis only. A comparison with the occurrence of a subdivision of EsD ² in Caucasians (EsD ⁵) suggests a remarkable difference in allele distribution of esterase D among races. Quantitative analysis showed a relatively low value of enzyme activity for this new allele. It is therefore emphasized that in addition to conventional electrophoresis, enzyme assay and further detection by isoelectric focusing are essential in analyzing the esterase D system.     

Esterase D polymorphism in Chinese and Japanese

Summary Esterase D phenotypes have been determined in Chinese and Japanese populations in the San Francisco area. The EsD¹ gene frequencies were 0.612 for the Chinese population and 0.582 for the Japanese population.This is communication 189 from the Forensic Science Group, University of California, Berkeley, USA     

Atypical segregation of esterase D: Evidence of a rare “silent” allele EsD 0

Summary Electrophoretic study of esterase D in 1027 mother-child pairs showed an atypical segregation of EsD alleles in one pair. The family analysis confirmed the evidence of a silent gene (EsD ⁰), which was observed in child, mother and grandfather. R banding of the metaphasal chromosomes revealed the normal appearance of the No. 13 pair, and no deletion of homologues No. 13 was observed in this family.     

Restriction and persistence of polymorphisms of HLA and other blood genetic traits in the Parakanã Indians of Brazil

Results concerning HLA types and 22 other blood genetic systems are reported for the Parakanã Indians of northern Brazil, a tribe that is notable for the light color and pilosity of some of its members. No clear evidence of Caucasoid admixture was found, but the Parkanã show unusual frequencies in the EsD¹, PGM¹₁, Gc², Cp^B, Fy^a, Di^a, and L^M genetic markers. In addition, the very rare Rh allele r^y is present, as well as what seems to be a new PGM₂ variant. There is very limited heterogeneity in the HLA system. All these distinctive features may have arisen through a combination of founder effects and genetic drift. However, low F_is values, as well as higher mean ages in heterozygous as compared to homozygous persons, suggest that an heterotic effect is counteracting these dispersive forces.     

Intra and intertribal genetic variation within a linguistic group: The Ge-speaking Indians of Brazil

A total of 562 individuals living in four villages of two Brazilian Indian tribes (Cayapo and Krahó) was studied in relation to blood groups ABO, MNSs, P, Rh, Lewis, Duffy, Kidd and Diego; haptoglobin, Gc, acid phosphatase and phosphoglucomutase types. These results were compared with those obtained previously among the Xavante, and the inhabitants of three other Cayapo villages, all of whom speak Ge languages; the ranges in gene frequencies observed in a representative series of South American Indians from all over the continent were also compiled. The Ge Indians are characterized by low frequencies ofR^z, medium frequencies ofR¹,R², R⁰, orr,Jk^a andPGM¹₁, and high frequencies ofGc² andACP^A when compared with other South American tribes. Genetic distance analyses based on six loci indicate that the intratribal variability observed among Cayapo is of the same order of magnitude as those obtained among the Xavante and Krahó, being much less pronounced than those observed among the Yanomama and Makiritare. The intertribal differences within this linguistic group are much less pronounced than those encountered among tribes that speak more differentiated languages.     

Genetic studies on free-ranging macaques of Cay Santiago. II. 6-phosphogluconate dehydrogenase and NADH-methemoglobin reductase (NADH-diaphorase).

For the population of 395 semi-free-ranging rhesus macaques (Macaca mulatta) that inhabited Cayo Santiago in 1976, 6-phosphogluconate dehydrogenase phenotypes of 378 animals were determined. Three phenotypes, controlled by two autosomal codominant alleles,PGD^A andPGD^B, were found by electrophoretic methods. The frequencies of the alleles are 0.898 and 0.102, respectively. The population, composed of five troops and peripheral males, is in Hardy-Weinberg equilibrium at this locus. The allele frequencies at the 6-phosphogluconate dehydrogenase locus in the population in 1976 were compared with frequencies in 1973; a statistically significant difference was found in one troop. The phenotypes of NADH-methemoglobin reductase (NADH-diaphorase) were determined electrophoretically for 372 animals. These phenotypes are probably the products of two autosomal codominant alleles,Dia¹ andDia², with frequencies of 0.786 and 0.214, respectively. The population is in equilibrium at this locus also. Tests of homogeneity at the dehydrogenase and reductase loci indicate that the allele frequencies are significantly different among the five troops in the population. Observed and expected phenotypic ratios in progeny were compared at the dehydrogenase and the reductase loci. The only significant deviation from expectation occurs among offspring of mothers heterozygous at the reductase locus. The observed distributions of alleles at the 6-phosphogluconate dehydrogenase locus and the NADH-methemoglobin reductase locus are probably the results of stochastic processes.     

Genetics of glucose phosphate isomerase and phosphoglucomutase in Aedes albopictus (diptera: culicidae)

Summary Glucose phosphate isomerase (E.C. 5.3.1.9) and phosphoglucomutase (E.C. 2.7.5.1) were found to be polymorphic in a laboratory colony of Aedes albopictus. The glucose phosphate isomerase locus is represented by two alleles resulting in three genotypes, while the phosphoglucomutase locus is represented by at least five alleles giving rise to a total of 15 genotypes. The inheritance of these two enzymes is of the Mendelian type with codominant alleles. Present data indicate that these genes are not linked.Of 105 mosquitoes analysed for these two gene-enzyme systems, the frequencies for glucose phosphate isomerase alleles are Gpi ^S=0.68 and Gpi ^F=0.32, while the frequencies for phosphoglucomutase alleles are Pgm ^A=0.16, Pgm ^B=0.11, Pgm ^C=0.19, Pgm ^D=0.30 and Pgm ^F= 0.24. The frequencies of the three glucose phosphate isomerase genotypes are in accord with Hardy-Weinberg expectations (X ₁ ² =2.74). Similarly, the frequencies of the 15 phosphoglucomutase genotypes probably do not differ significantly from Hardy-Weinberg expectations (X ₁₀ ² = 18.45).     

A long-term study on interactions between the Adh and αGpdh allozyme polymorphisms and the chromosomal inversion In(2L)t in a seminatural population of D. melanogaster

The Adh and αGpdh allozyme loci (both located on the second chromosome) showed considerable fluctuations in allele frequencies in a seminatural population of Drosophila melanogaster during 1972–97. Both long-term and short-term fluctuations were observed. The short-term fluctuations occurred within almost all years and comparison of allele frequencies between winters and summers showed significantly higher Adh^S (P < 0.001) and αGpdh^F (P < 0.01) allele frequencies in summers. Frequencies of these alleles were significantly positively correlated with environmental temperature, suggesting the adaptive significance of these allozyme polymorphisms. Frequency changes of the Odh locus (located on the third chromosome) showed no seasonal pattern and were not correlated with environmental temperature. Almost all short-term and long-term increases in Adh^S frequency were accompanied by a corresponding decrease in αGpdh^S frequency (r = –0.82, P < 0.001) and vice versa. Further analysis showed that gametic disequilibria between the Adh and αGpdh loci, which frequently occurred, were due to the presence of inversion In(2L)t located on the same chromosome arm and In(2L)t frequencies were positively correlated with environmental temperature. Gametic disequilibria between Adh and Odh and between Odh and αGpdh were hardly observed. Because In(2L)t is exclusively associated with the Adh^S/αGpdh^F allele combination, the observed correlated response in Adh/αGpdh allele frequencies is (at least partly) explained by hitchhiking effects with In(2L)t. This means that the adaptive value of the allozyme polymorphisms has been overestimated by ignoring In(2L)t polymorphism. Fluctuations in Adh allele frequencies are fully explained by selection on In(2L)t polymorphism, whereas we have shown that αGpdh frequency fluctuations are only partly explained by chromosomal hitchhiking, indicating the presence of selective differences among αGpdh genotypes in relation with temperature and independent of In(2L)t. Frequency fluctuations of αGpdh and In(2L)t are consistent with their latitudinal distributions, assuming that temperature is the main environmental factor varying with latitude that causes directly or indirectly these frequency distributions. However, the results of the tropical greenhouse population show no correlation of Adh (independent of In(2L)t) and Odh allele frequencies with environmental temperature, which may indicate that the latitudinal distribution in allele frequencies for these loci is not the result of selection on the F/S polymorphism in a direct way.     

Glucose phosphate isomerase and 6-phosphogluconate dehydrogenase polymorphism in Malaysian leaf monkeys

Glucose phosphate isomerase and 6-phosphogluconate dehydrogenase were found to be polymorphic in Malaysian leaf monkeys. Two glucose phosphate isomerase electrophoretic phenotypes were presumed to be homozygous. Three 6-phosphogluconate alleles and four electrophoretic phenotypes were present. The allele frequencies inPresbytis obscura werePgd ^A=0.64,Pgd ^B=0.27 andPgd ^C=0.09. The frequencies of the 6-PGD phenotypes inP. obscura were not in accord with Hardy-Weinberg expectations. All the biochemical markers examined show identical electrophoretic patterns in the Malaysian leaf monkeys.     

Distribution of hereditary blood groups among Indians in South America. II. In Peru

Blood specimens were procured from the following putatively pure Indians of the Peruvian rain forest: 90 Piro and 89 Campa on the Urubamba and Tambo rivers, 142 Shipibo and 14 Isconahua on the Rio Ucayali near Yarinacocha, 151 Aguaruna at Santa Maria de Nieva, where the Marañon and Nieva rivers join, and from 122 Ticuna and 9 Yagua near the Brazilian border on the Amazon. Specimens from highland Indians were obtained from 93 Aymará and 181 Quechua at Puno and environs. These 891 specimens were tested for antigens in the A-B-O, M-N-S-s, P, Rh-Hr, Lutheran, K-k, Lewis, Duffy, Kidd, and Diego (Di^a) systems, and for the Wright (Wr^a) aglutinogen. Serum samples from these bloods were tested for haptoglobins and transferrins and hemolysates were prepared and examined for hemoglobin types. Results for these tests with claculated gene frequencies are presented, for the most part, on appropriate tables. A map is included to show the locations of the populations from which blood samples were procured. As in South American Indians generally, frequencies are high for the O gene it being the only gene of the ABO system which appears in isolated jungle populations and the Aymará. Gene frequencies are usually high also for M, s, R¹ (CDe), R² (cDE), Lu^b, k, Le^H, and Fy^a; and low or absent for A, B, N, S, Mi^a, Vw, R^o (cDe), r (cde), Lu^a, K, Le¹, Fy^b, and Wr^a. The Diego (Di^a) gene is present but varies greatly in frequencies among tribes. Hp¹ gene frequencies vary from 0.44 to 0.69 among the Peruvian Indians tested. Transferrin CD was encountered in only one population i.e., in 3 of 86 Piro (gene frequency Tf^D= 0.02). All others were C. All Peruvian Indian bloods tested electrophoretically contained only hemoglobin (A) as a major component.     

Distribution of hereditary blood groups among Indians in South America. 3. In Bolivia

Blood samples were procured from the following populations of putatively pure Indians in Bolivia: 503 Aymará from the Altiplano and Yungas, 30 Chama, 11 Tacana, 14 Chácobo, 109 Itonama, 67 Moré, and 27 Sirionó from the Beni and lowland rainforest. Erythrocytes from these 761 specimens were tested for antigens in the A-B-O, M-N-S-s, P, Rh-Hr, Lutheran, Kell-Cellano, Lewis, Duffy, Kidd, and Diego systems, and for the Wright agglutinogen. The serum samples were tested for haptoglobins and transferrins; and hemolysates were prepared and examined for hemoglobin types. Results of these tests are presented as phenotypes and calculated gene frequencies on appropriate tables. A map is included to show the locations of the populations from which blood samples were obtained. Frequencies are generally high for the O gene, it being the only gene of the ABO system which appears in the Chama, Chácobo and Sirionó. The presence of A₁, A₂ or B genes in the Bolivian Indians is interpreted as being most probably of caucasoid introduction. Excepting the Sirionó the frequencies are high for M and low for N genes as is usual for Amerinds, the M gene being the only one detected in the Chama. The s gene frequency in high and the S low except in the small isolated Chácobo population in which S gene frequency is extremely high for Amerinds. Inbreeding and perhaps genetic drift in this small isolate may account for this aberrancy from normal. The Bolivian specimens presented the high frequencies for genes R¹ (CDe) and R² (cDE) and the low frequencies for genes r (cde) and R⁰ (cDe) usually observed in American Indians. The Lu^a factor was observed in only one of 120 Aymará at Santa Fe in the Yungas. The Lu^a factor, when observed in Amerinds, suggests foreign introduction of the responsible gene. Fy^a gene frequencies are consistently high and excepting the Aymará and Chama so also are Jk^a frequencies. Frequencies for the Diego (Di^a) factor vary from 3.70% in 27 Sirionó to 73.33% in 30 Chama. No K, Mi^a, Vw or Wr^a antigens were demonstrable in the Indian blood samples from Bolivia. Phenotypes and calculated gene frequencies for haptoglobins and transferrins are presented. All Bolivian Indian bloods tested electrophoretically contained only hemoglobin (A) as a major component.     

Distribution of hereditary blood groups among Indians in South America. V. In northern Brazil

This paper reports the results of tests made for hereditary antigens in blood samples procured from Indians in northern Brazil. Specimens were procured from 423 putatively full-blood persons of the following tribes: in the province of Roraima from 261 Macuxi, 48 Uaica, 27 Xirixano, 10 Uapixana, 9 Cacarapai and 9 Paramiteri; in Pará from 21 Assurini; and in Amapá from 38 Galibi. Erythrocyte samples were tested for factors in the A-B-O, M-N-S-s, P, Rh-Hr, Lutheran, Kell-Cellano, Lewis, Duffy, Kidd and Diego systems. Serum samples were tested for haptoglobins and transferrins. Hemolysates, prepared from whole blood, were tested for hemoglobin types. The results are presented on appropriate tables as number and per cent of phenotypes for the various blood group anigens and their calculated gene frequencies. Locations from which blood samples were procured are listed in the tables and shown on a map (fig. 1). All the 423 samples except one Macuxi belonged to group O. The Uaica tribe had a low frequency for M (0.534). All others showed the high frequency usually observed in Amerinds. The s allele was high in all except the Galibi in which the frequency was (0.500). Frequencies for P² was higher than for P¹ in all except the Assurini and Galibi, theirs was high for P¹ (1.00) and low for P² (0.00). The frequencies for R¹ (CDe) and R² (cDE) were high and all others in the Rh-Hr system were low or absent. All specimens were positive for Cellano (k) and negative for Kell (K). There was a complete absence of Lewis (Le₁), excepting in the Uaica and Xirixano in which populations Fy^a allele frequencies were higher than 0.500. The distribution of the Jk (a+) phenotype and corresponding ellele frequencies varied widely in Brazilian Indians as did those for Diego (a+). The haptoglobin Hp¹ allele frequencies were in essential agreement with those reported elsewhere for Indians in South America, and all transferrins determined were classified as Tf C. All samples tested for homoglobin types contained homoglobin (A) as a major component, but five members of the Galibi tribe possessed hemoglobin (S) as well.     

Transfer of plasmids and chromosomal genes amongst subspecies ofBacillus thuringiensis

Summary The plasmids pBC16 and pC194 fromBacilus thuringiensis subsp.israelensis strains A084-16-194 were transferred to 25 subspecies ofB. thuringiensis by a conjugation-like process using broth mating technique. The frequencies of transfer varied considerably between different mating pairs, ranging from 1.1×10^–9 to 9.8×10^–5. Additionally, chromosomal transfer could also be demonstrated in tenB. thuringiensis subspecies with very low frequencies (4.3×10^–9 to 3.7×10^–7). The intersubspecies matings within a group of eight subspecies strains gave higher frequencies of transfer than the matings between the subspecies. Furthermore, the results indicated that the capability to transfer plasmids among these various subspecies did not depend on the presence of large plasmid.     

Distribution of blood groups among indians in South America. VI. In Paraguay

This paper on the distribution of hereditary factors in the blood of Indians in South America, reports the results of tests made on samples procured from Paraguayan Indians. Specimens from putatively full-blood persons were obtained from the following tribes: 88 Chamacoco, 36 Moro, 85 Chulupi, 207 Lengua, 100 Toba, 20 Yam Lengua, and 51 Guayaki, These 587 Samples were tested for factors in the A-B-O, M-N-S-s, P. Rh-Hr, Lutheran, Kell-Cellano, Lewis, Duffy, Kidd, and Diego systems. Serum samples were tested for haptoglobins and transferrins. He molysates, prepared from whole blood, were tested for hemoglobin types. The results are presented on appropriate tables as number and per cent of phenotypes for the various blood group antigens and their calculated allele frequencies. Locations of the populations from which blood samples were procured are listed on the tables and shown on a map (fig. 1). Of the 587 samples all except two Chamacoco belonged to group O. High frequencies are reported generally for M, s, P, R¹ (CDe), R² (cDE), k (100%) and Fy alleles in Paraguayn Indians. Low frequencies were generally reported for N, S, r (cde) and R° (cDe) alleles. There was a wide variation in frequencies for Di, Jk, and haptoglobin Hp¹. All tested for transferrins were classified as Tf C and all contained hemoglobin (A) as a major component. The following antigens were completely absent: Mi^a, Vw, p, P^k, r^y (CdE), K, and Le¹. Most notable is the unusual distribution of hereditary blood antigens among the Guayaki and Moro. The Guayaki had 100% P₁ and Fy^a; they were higher in R° (cDe), R¹ (CDe), and Jk^a; and lower in R² (cDE) and Hp¹ genes than other Indians; and Di was absent. The Guayaki differed from the other Indians also in having fair skin. The Moro were lower in the P¹ and Jk gene frequencies than is usually found in Amerinds, and the Di gene was absent. The Chamacoco also had an exceptionally low frequency for the P¹ gene (0.261).     

Distribution of hereditary blood groups among Indians in South America. IV. In Chile with inferences concerning genetic connections between Polynesia and America

This is the fourth paper in a series on the distribution of blood groups among Indians of South America. It reports the findings on the Indians of Chile and the Polynesians of Chile's Easter Island. Blood specimens were procured from the following putatively pure Indians and unmixed Polynesians: 44 Alacaluf of Puerto Eden, Isla Wellington, 141 Mapuche (Araucanian) of Lonquimay, Malleco Province, 80 Atacameños of Antofagasta Province, and 45 Polynesians of Easter Island. These 310 samples were tested for blood factors in the A-B-O, M-N-S-s, P, Rh-Hr, Lutheran, K-k, Lewis, Duffy, Kidd and Diego systems, and for the Wright (Wr^a) agglutinogen. Serum samples were tested for haptoglobins and transferrins. Hemolysates prepared from the blood clots were tested for hemoglobin types. The results are presented as phenotype incidences and calculated gene frequencies in appropriate tables. Locations of the populations from which blood samples were procured are shown on two maps. The high frequencies for the O gene usually reported for South American Indians obtain in putatively pure Chilean Indians but A¹ is high in Easter Island Polynesians. In both Indians and Polynesians M, s, R¹ (CDe), R² (cDE), Lu^b, k, Le^H, and Fy^a gene frequencies are high and B, N, S, Mi^a, Vw, Rº (cDe), r (cde), Lu^a, K, Le¹, Fy^b, and Wr^a (Ca) are low or absent. The Diego (Di) gene is present in the Mapuche and Atacameños but absent in the Alacaluf and Polynesians. Hp¹ gene frequencies were determined only in the Alacaluf and Atacameños, in which they are 0.48 and 0.67 respectively. Transferrins were determined for the Alacaluf and Atacameños Indians and all were classified as Tf C. All Chilean Indian and Polynesian specimens were tested electrophoretically for hemoglobin types and all contained only hemoglobin (A) as a major component.     

Molecular cloning of theEnvC gene ofEscherichia coli

DNA fragments complementing theenvC mutation could be isolated by cloning chromosomal DNA in the vector pUH84. When the frequencies of transformation and the frequencies of restoring theenvC ⁺ phenotype were compared, the high copy number hybrid plasmids complemented with a frequency of 10^–5. After subcloning theenvC-complementing DNA fragment into the low copy number plasmid pLG339, efficient complementation was achieved by spontaneous integration of the IS2 element ofEscherichia coli. By nucleotide sequence analysis, a potential promoter, a ribosome-binding site, and an unidentified reading frame were detected in the respective DNA fragment.     

Hemoglobins, haptoglobins, and transferrins in indigenous populations of Kenya

Hemoglobin, haptoglobin, and transferrin phenotypes were determined by means of starch-gel electrophoresis for a sample of 226 Kenya hospital patients. Allele frequencies were: Hb_β^S=0.081; Hp¹=0.057; Tf^D=0.038. Hemoglobin S was the only aberrant hemoglobin found in this sample. Transferrins C and D were the only transferrins found. Hemoglobin and transferrin phenotypes were also determined for a sample of 201 newborn Kenya infants. One of these infants had hemoglobins F, S, and C, eight had hemoglobins A, F, and S, and the remainder had hemoglobins A and F. Transferrins C, B, and D were found in this sample. Allele frequencies were: Tf^B=0.008; Tf^D=0.019.     

Distribution of hereditary factors in the blood of Indians of the Gila River, Arizona

This paper reports the distribution of blood groups, A-B-H secretors, haptoglobins, transferrins and hemoglobin types among Indians of the Gila River Valley in Arizona. Specimens were procured from the following putative full-bloods: 909 Pima, 37 Papago, and 124 Maricopa; and from the following known mixed-bloods: Pima-Papago 134, Pima-Maricopa 26, Pima-Other Indian 41, Pima-Caucasian 33. These 1304 samples were tested for factors in the A-B-O, M-N-S-s, P, Rh-Hr, Lutheran, Kell-Cellano, Lewis, Duffy, Kidd and Diego blood group systems, and for additional blood factors (Wr^a), Do^a, Vel, Yt^a, Co^a, Gy^a, Sav, and L. W. Serum samples were tested for haptoglobins and transferrins. Hemolysates, prepared from whole blood, were tested for hemoglobin types. The results are presented on appropriate tables as number and per cent of phenotypes for the various blood group antigens and their calculated allele frequencies. Locations of the populations from which blood samples were procured are shown on a map (fig. 1). Tests made by earlier workers on the blood of Arizona Indians and related tribes are presented for comparison and discussed. The usual high frequencies for allele O reported in Amerinds was found among the putatively full-blood Gila Indians; the 124 Maricopa presented the maximum frequency of 1.000. High frequencies were reported generally for M, s, P¹, R¹ (CDe), R² (cDE), k (100%) Fy, and Do^a alleles. Low frequencies were reported for N, S, r (cde), R° (cDe), fy, Le¹w and Di^a (Pima only). There was a wide variation in frequencies for jk, and Hp¹, and there were 17 Transferrin Tf B₁C observed in 270 Pima samples tested. All the remaining were classified as Tf C except two Tf B;C from mixed-bloods. All samples tested for Vel, Yt^a, Co^a, Sav, and Hemoglobin (A) showed the maximum frequency (1.000) for their genes. The following antigens were completely absent: Lu^a, Mi^a, Vw, Mt^a, p, P^k, r^y (CdE), K, and Wr^a. The results of this study suggests that the Papago tribe presents fewer genes of non-Indian origin than the Pima, and the Maricopa least of the three populations.     

The geographic distribution of flavone-glycosylating genes in Silene pratensis (Rafn.) Godron and Gren. (Caryophyllaceae)

In Silene pratensis three loci (g, gl and fg) control the glycosylation of isovitexin. Three alleles are known for both the g-locus (g, g ^G and g ^X ) and the gl-locus (gl, gl ^A and gl ^R ); for the fg-locus there are only two alleles (fg and Fg). The distribution of these alleles over 285 European populations of S. pratensis has been investigated. It was concluded that there are three different chemical races within S. pratensis in Europe. The first race contains the populations in western and southern Europe and displays high frequencies of g ^G , gl and fg. The frequencies of g ^G and gl ^R are very high in the second chemical race, which can be found in the USSR, Scandinavia and eastern Poland. The third chemical race occurs in central Europe and in this race the frequencies of both g and gl ^R are high, Fg has low to moderate frequencies in the second and third groups. The alleles gl ^A and g ^X are seldom found in S. pratensis, but are present in the closely related S. dioica. They do occur with low frequencies in some populations of S. pratensis, possibly as a result of hybridization with S. dioica.