Human genetic variation in the Sierra de Gredos mountain (Central ...

ANNALS OF HUMAN BIOLOGY, 1996, VOL. 23, NO. 3, 213--221

Human genetic variation in the Sierra de Gredos mountain (Central Spain): study of several polymorphisms P. MORAL~, T. SANDIUMENGE~, S. VIVES'~, N. LUTKEN~, F. ORTEGA~, M. D. MARRODAN§ and V. FUSTER§ tFacultad de Biologia, Universidad de Barcelona SResidencia Sanitaria de la Seguridad Social Nuestra Sefiora de Sonsoles, Avila §Facultad de Biologia, Universidad Complutense, Madrid

Ann Hum Biol Downloaded from informahealthcare.com by Dr V. Fuster on 10/23/14 For personal use only.

Received 20 March 1993; revised 15 December 1995

Summary. The variation in the third component of human complement (C3) and orosomucoid (ORM1) serum proteins, and 6-phosphogluconate dehydrogenase (6PGD), adenosine deaminase (ADA), esterase D (ESD), and acid phosphatase (ACP1) red cell enzymes was examined in two samples from autochthonous populations living on either side of the Sierra de Gredos range in Central Spain. These results support only a moderate differentiation between Gredos subpopulations, which is discussed in relation to other Iberian Peninsula groups. Allele frequencies in Gredos show a remarkable heterogeneity as contrasted with other Iberian populations for most of the markers examined; relatively high C3"S, ORMI*F, ADA*I, ESD*2, and ACPI*C gene frequencies are characteristics of Gredos samples. This differentiation was more marked for the Northern Gredos population and could be related to the geographical peculiarities of this region.

1. Introduction The study of populations living in mountain regions has a particular interest in anthropology. The topography of those areas may constitute a certain barrier to the free diffusion of people and genes, and may sometimes be responsible for marked differences in gene frequencies and other anthropological characteristics between neighbouring population groups. In this context the present paper deals with the variation of several genetic markers in two populations settled in a mountainous area of Central Spain: the Sierra de Gredos. This mountain chain, located in the province of Avila (figure 1), forms part of the Sistema Central, the biggest mountainous range in Central Spain, which divides the Castilian Plateau, running S W - N E . There are geographical and climatic differences between the two slopes of the Sierra de Gredos which have influenced socioeconomic differences among its populations, devoted mainly to cattle raising and forestry on the northern side and to agriculture on the south. The population living on the northern part (a total of 35,688 according to the 1981 census) is distributed in 38 small villages scattered along the narrow valleys of the Tormes and Alberche rivers bounded by high mountains, with an extreme climate. The southern slope, where the climatic conditions are less extreme, opens to the Southern Castilian Meseta and its population (34,245 inhabitants) is concentrated in 22 villages (with sizes from 367 to 6604 inhabitants) distributed along the Ti6tar river. Historical evidence of human presence in the region goes back to 1000 BC tO the Vettones, a shepherding people probably of Celtic origin, and later to the Romans and Visigoths in the sixth century. After a general depopulation during the first centuries of Moslem occupation due to its position as a frontier land with frequent wars, a slow repopulation from neighbouring areas took place throughout the Middle 0301-4460/96 $12"00 (c) 1996 Taylor & Francis Ltd.

214

P. Moral et al. 8°

W

4" W

O"

W

04


42° N

5° W

40 °

N

38 °

N

4 ° 30"W

Nor:hem Plateau

40 ° 30"N

.40" N

Southern

Pla

Figure 1. Geographical location of the Sierra de Gredos region and of the populations used for comparisons: (1) Northern Gredos (Tormes-Alberche valley), GRN; (2) Southern Gredos (Ti6tar valley), GRS; (3) Vizcaya, VIZ; (4) Galicia, GAL; (5) Northern Portugal, PON; (6) Pallars Sobirfi~ valley, PAL; (7) La Garrotxa region, GER; (8) Barcelona, BAR; (9) Eastern Andalusia, ANE.

Ages, with possibly small contributions from ethnic minorities such as Jews, Moors, and Mozarabs seeking asylum in the mountains of the Sierra de Gredos (Troitifio 1990). In recent times, as with many other Spanish rural areas, intense emigration to industrialized regions has caused a marked fall in population sizes since 1960. The particular geographical characteristics of this region have led us to an anthropological study in order to assess the possible influence of the Sierra de Gredos on population differentiation. The starting point was that if this mountain chain-reaching altitudes of 2600 m and crossed by only three natural passes had acted as a barrier to population movement, especially during winter, then biological differences between the populations from the two sides could be expected. In this respect, previous analysis of marriages in this region (Fuster, Martin, Mesa and Marrodan 1993) indicates that the marital mobility between the two slopes of the Sierra de Gredos has been less than 2~o during the last 100 years. Furthermore, results on blood groups show a certain degree of genetic differentiation between northern and southern Gredos populations (Mesa, Martin, Fuster and Fisac 1994). Here we present

Genetic polymorphisms in Gredos, Central Spain

215

new genetic data on six classical markers yielding additional genetic information about the populations from either side of Gredos mountain. The degree of genetic heterogeneity between Gredos subpopulations is discussed in relation to other Iberian Penninsula groups.


2.

Material and methods Blood samples from healthy unrelated adults of both sexes were obtained in collaboration with the Haematology and Haemotherapy Service of Nuestra Sefiora de Sonsoles Hospital in Avila, during blood collection campaigns carried out in different villages scattered throughout the three main valleys of the Sierra de Gredos: Tormes, Alberche, and Ti6tar (figure 1). In order to obtain a sample representing the autochthonous population, individuals were questioned and classified according to the place of origin of their ancestors. Only donors whose four grandparents were natives of the same valley were considered in this study. The total sample, made up of 202 subjects, was subdivided into subsamples called Northern Gredos and Southern Gredos. The former includes persons from the two valleys of the northern slope (Tormes and Alberche) due to the geographical continuity of these areas and to the fact that numbers from each valley were too small, while the latter group contains only individuals originating from the Ti6tar valley. Sera were examined for the third component of human complement (C3) and for orosomucoid (ORM1) types. C3 genetic variants were detected by horizontal electrophoresis on agarose gels, as described by Teisberg (1970) with minor modifications. ORM1 phenotypes were identified by isoelectric focusing (IEF) in polyacrylamide gels (C = 5~,, and T = 3~) using a pH 2.5-4 and pH 2.5-5 ampholine mixture, followed by silver staining. From red blood cells the variants of 6-phosphogluconate dehydrogenase (6PGD; EC 1.t.1.44), adenosine deaminase (ADA; EC 3.5.4.4), esterase D (ESD; EC 3.1.1.1), and acid phosphatase (ACPI; EC 3.1.3.2) were determined. Separation and staining of the isoenzymes were carried out according to the protocols by Harris and Hopkinson (1976), by means of horizontal electrophoresis on starch gels. ESD*5 gene products were typed by 1EF at pH 4 6, following the technique of Divall (1984). Allele frequencies were calculated by direct gene counting, and the Hardy-Weinberg equilibrium was tested by chi-square. Genetic differentiation between populations was evaluated through chi-square tests and FsT. Genetic distances and principalcomponent analysis were also performed as a measure of the interpopulation variation. 3.

Results and discussion Phenotype and allele frequencies for the six genetic polymorphisms examined in the two Gredos populations are presented in table 1. When possible, the validity of Hardy Weinberg equilibrium was evaluated by chi-square, and the genotype distributions observed showed good agreement with the expected Hardy Weinberg proportions. A rare variant C3"So. 4 was identified in the Northern Gredos subpopulation. Slight gene frequency differences between the two Gredos subsamples are especially visible for ORMI*F and ACPI*B alleles, with higher values in the Northern group. However, these differences were not statistically significant either for individual markers (C3: X2 = 1.34, p--0.513; ORMI' Z~ = 2-34, p = 0-126; PGD: Z~ = 0.39,

216

P. Moral et al. Table 1.

Phenotype and allele frequency distribution in the Sierra de Gredos valleys.

Phenotype

Northern Gredos Observed

C3 S FS F SS04


Total ORMI F FS S

Total PGD A AC C

Total ADA 1 21 2

Total ESD 1 21 2 51 52

Total ACP1 A AB

B AC BC C

Total

Expected

Southern Gredos

Gene frequencies

77 19 4 1 101

74-94 23.26 1.80 0.86

48 46 11 105

48-01 45.98 I1.01

96 7 -103

96.12 6.76 0.12

PGD*A PGD*C

0.966_+0.013 0.034_+ 0.013

104 I

104.00 0.99 0.01

ADA*I ADA*2

0-995 _+ 0-004 0.005 + 0.004

--

C3"S C3*F C3"S0.4 (Z12= 3.48,

0.861 _+ 0.024 0.134 _+ 0.024 0.005 + 0.004 p = 0.062)t

ORM*F 0.676+_0.032 ORM*S 0.324+_0.032 (Z2 < 0.01, p = 0.997)

105

Observed

Expected

Gene frequencies

63 22 3

62.23 23.54 2.23 0-00

C3"S C3*F

88

0.841 + 0.026 0.159 _+ 0-026

(Z 2 = 0.38, p = 0.538)

38 41 18 97

35-28 46.44 15.28

79 6 1 86

78.19 7.63 0-18

PGD*A PGD*C

0.953 Jr 0.016 0.047_+ 0.016

79 4

79-05 3.90 0.50

ADA* 1 ADA*2

0.976 _+ 0.011 0.024 _+ 0.011

ESD*I ESD*2 ESD*5

0.851 + 0.025 0.139 + 0.024 0.010 ___0.007

ORM*F 0.603 + 0 . 0 3 5 ORM*S 0.397+0.035 (Z 2 = 1.33, p = 0-249)

83

72 28 3 2 1 106

71.41 28.73 2-90 2.46 0"50

8 39 41 7 9 1 105

9-15 38"38 40'24 5-31 11"14 0-77

ESD*I ESD*2 ESF*5

0.821 _+ 0-026 0.165 + 0.025 0-014 _+ 0.008

(Z 2 = 0'02, p = 0"890)~-

ACP*A ACP*B ACP*C

0-295 +__0"031 0-619 _+ 0-033 0.086_+0.019

(Z32= 1.18, p = 0.757)

69 25 i 2 -97

70.17 22.96 1.88 1.70 0-26

9 37 31 8 11 1 97

10"23 35'72 31-19 6"82 11.91 1-14

(z~ = 0-60, p = 0440)~

ACP*A ACP*B ACP*C

0-325 + 0"033 0.567 +_ 0.035 0-108+0-023

0~32= 0.484, p = 0-922)

t C3 SSo.,, a n d E S D 5 phenotypes not included in Hardy Weinberg equilibrium calculation.

p = 0"534; ADA: Z2 = 2.64, p = 0"104; ESD: Z22 = 0-67, p = 0"714; ACPI: Z2 = 1.27, p = 0-529) or for chi-square pooled over all loci (Z 2 = 8.65, p = 0.470). In comparison with other population data from the Iberian Peninsula, C3*F gene frequencies in the two Gredos populations are particularly low (Peninsula values range from 0-164 to 0-277). This situation is especially evident in the case of the Northern Gredos subsample, where C3*F frequency is significantly lower than most Iberian groups (13 out of 16 populations compared) including two samples from the neighbouring population of the city of Madrid (Goedde, Hirth, Benkmann, Pellicer, Pellicer, Stahn and Singh 1973, Regueiro and Arnaiz-Villena 1984). For the ORM1 system the Northern Gredos group also presents the lowest O R M I * S till n o w found in Iberian Peninsula (with a range of 0.369-0.460 over a


Genetic polymorphisms in Gredos, Central Spain

217

total of eight population data reviewed), showing significant differences with the values reported in Galicia, Basque Country, and Northern Portugal (Montiel, Carracedo, Blazquez-Caeiro and Andrade-Vide 1990). PGD*C allele frequencies in Gredos are similar to the highest values in the Iberian range (0-000 0.047, from a total of 30 data reviewed), and particularly different from geographically diverse groups such as Basques (Aguirre, Vicario, Maz6n, Estomba, Pancorbo, Arrieta, P6rez-Elortondo and Lostao 1991), Cadiz (Southern Spain; Castellano and Martinez-Jarreta 1991), Northern Portugal (Dos Santos 1983), and Madrid (Goedde et al. 1973). As for the ADA system, low ADA*2 allele frequencies seem to characterize Gredos populations inside the Iberian variation (0-007 0"080, over a total of 25 sample data reviewed). These frequencies, especially in the case of Northern Gredos subsample, are similar to the lowest Peninsula values reported among Basques (Aguirre, Vicario, Mazdn, Estomba, Pancorbo, Arrieta, Perez-Elortondo and Lostao 1991; Manzano 1991) and clearly different from other Peninsula areas including the nearby population of Madrid (Goedde, Herth, Benkmann, Pellicer, Pellicer, Stahn and Singh 1973). The Gredos populations show relatively high ESD*2 gene frequencies in relation to the Iberian variation (0.034 0.174, from a total of 36 population data) and are particularly different from Basque groups, among which the lowest values have been found (Aguirre et al. 1991). ESD*5 gene frequencies are similar to those described in other Peninsula areas (0-007 0-024 range from 11 population data). For the ACP1 system the most relevant characteristics of Gredos populations are high frequencies of ACPI*C gene in contrast to an Iberian average value of 0.035 (0.000 0.121 range, out of 46 sample data), and relatively low ACPI*B frequencies (Peninsula range 0.558 0.796). These results are significantly different from those of Basques (Aguirre et al. 1991) and of other population samples from Central Spain (Pancorbo, Maz6n and Lostao 1986, Gremo 1988), and do not support any clinal distribution in the Iberian Peninsula. In an attempt to summarize the global genetic position of the Gredos samples we obtained joint information on the set of markers studied in other Iberian Peninsula populations, and data were available for seven groups, some of which correspond to geographically well-limited units comparable with that of Gredos: Galicia (GAL) and Northern Portugal (PON) in the northwest (Goedde et al. 1973, Montiel et al. 1990, Carracedo and Concheiro 1983, Dos Santos 1983, Amorim and Siebert 1982, Amorim, Rocha and Santos 1994) Vizcaya (VIZ) from Spanish Basque Country (Goedde et al. 1973, Montiel et al. 1990, Aguirre et al. 1991), Pallars Sobir5 valley (PAL) in Eastern Pyrenees (Moral, Panadero and Moreno 1986, Panadero, Moreno and Moral 1986, Sandiumenge, Vives and Moral 1993), the Garrotxa pre-Pyrenean region (GER) in Gerona province (Moreno and Moral 1983, Sandiumenge et al. 1993), Barcelona (BAR) in the northeast (Panadero 1988, Moreno and Moral 1981, Moral and Panadero 1984), and Eastern Andalusia (ANE) in the south (Castellano and Martinez-Jaretta 1991). Their geographic locations are illustrated in figure 1. On the basis of gene frequencies, chi-square for heterogeneity between populations and FsT (table 2) show a remarkable interpopulation genetic differentiation for PDG, ADA, ESD, and ACP1 polymorphisms and for the average across all loci. A genetic distance analysis (table 3) using the co-ancestry coefficient of Reynolds, Weir and Cockerham (1983) indicates that the distance between Gredos subpopulations is double those among the rest of non-Basque Iberian groups (average

P. Moral et al.

218

Table 2. Chi-square and FST for heterogeneity values.

System

~2

d.f.

FST

Z~(1)2

d.f.

C3 ORMI

26.871 8-77 48.63*** 24.30*** 63.17"** 67.64*** 239.37***

16 8 8 8 8 16 64

0-002 0.000 0.004 0.005 0-008 0.003 0-003

21.23 0.00 26-06*** 32.42*** 60.58*** 54.24*** 194.53"**

16 8 8 8 8 16 64

PGD ADA ESD ACP1

Average

1Determined by Workman and Niswander'smethod (1970).


*** p < 0.001.

Table 3.

GRN GRS VIZ GAL PON PAL GER BAR

Genetic distances ( x 10,000) between different Iberian populations.

GRS

VIZ

GAL

PON

PAL

GER

BAR

ANE

57

210 173

134 94 29

79 59 109 36

43 55 114 43 24

76 59 49 16 42 26

70 49 50 20 43 29 3

75 46 77 31 24 40 16 18

Population abbreviations defined in figure l.

distance: 27). This intra-Gredos genetic divergence is only slightly lower than both the Gredos vs. non-Basque Iberians average (70) and Basque vs. non-Gredos comparisons (average 71), but clearly lower than the highest (three to four times greater) differentiation found between Gredos and Basques (average 191). It is as if Gredos and Basques represented the extremes of Iberian variation, the rest showing an intermediate position. This situation can be made more explicit by measuring population genetic variation through principal-component analysis. Figure 2 shows for the first component scores plotted against those for the second component. These two factors explain 50"7~o and 18-4~o respectively (69-1~o accumulated) of the total variance. The population distribution due to the first component underlines the main separation between Gredos and Basques, and to a lesser extent from the rest of Iberian groups, the Pyrenean population of Pallars Sobirfi showing the greatest proximity with Gredos. The high correlations between the first component with the original frequencies of alleles ACP*C (r = 0'921), C3"S (r = 0.815), PGD*C (r = 0-721), and ESD*2 (r = 0'701) seem to indicate that, from the markers studied, high frequencies for these alleles are the most relevant traits of Gredos populations within the Iberian Peninsula variation. The second component, mainly correlated with ACP*A gene values (r = -0'735), contributes to the separation between Gredos subpopulations as well as among other Peninsula series. Concerning a possible heterogeneity within Gredos region, the results of this analysis only support a moderate interpopulation differentiation, which ranks with some of those between geographically distant Iberian groups (see table 3). Although chance cannot be ruled out, because of the lack of statistically significant differences for the six loci examined, our data are not, however, in contradiction with those

Genetic polymorphisms in Gredos, Central Spain 2nd Principal Component

219

118.4%)

GRN

VIZ PAL

P0N GAL BAR •


GER

"I ANE .

" GRS

-2 I

I

-2

-1

L 0

1

2

1st Principal Component (50.7%) Figure 2. Principal-component analysis: plot of the first and second components. Population abbreviations defined in figure 1. from blood groups (Mesa et al. 1994) and dermatoglyphics (Martin, Mesa, Fuster and Moral 1995), pointing to a certain degree of population divergence between the two slopes of the Sierra de Gredos, which could be related to the population history of this region and its geographical characteristics. Sharing the same ancient population substratum, both slopes probably received migration contributions from slightly different groups during the repopulation throughout the ninth and twelfth centuries AD, when the most relevant population movements took place in historical times. These small differences could have been preserved by the effect of the mountain range limiting the free gene flow between two sides, in accordance with a reduced marriage movement between the slopes (less than 2~o) and a relative excess of marital contribution from Northern and Southern Castilian Plateaus to the northern and southern slopes, respectively (Fuster et al. 1993). On the other hand, the results of the six polymorphisms analysed indicate a particular position of the Gredos populations within the Iberian Peninsula. In fact, the extreme values for several markers in comparison to the Iberian variation described so far, the genetic distance relationships, and the principal-component analysis might suggest a wider genetic variation in the Peninsula than that inferred from the available present data, probably in relation to sampling strategies, since many of them correspond to wide areas where the circulation is great and possible differences could have been diluted by recent migration. It is not possible to understand the history of the Gredos region, as in many other Spanish areas, without gene exchanges with surrounding populations. However, one must not neglect the possibility of an evolution in autochthonous populations under particular conditions, such as could have occurred in some Pyrenean groups (Fernet, Jacquart and Jakobi 1975). In this matter it is worth mentioning that the differentiation from other Iberian groups is more marked for the Northern Gredos subpopulation (showing extreme

220

P. M o r a l et al.

values for five out of the six markers) where isolation m a y have had a more significant effect, according to the geographical differences within the Sierra de G r e d o s region. These observations have only provisional value due to the low n u m b e r of markers considered; new genetic data a n d i n f o r m a t i o n from other sources are required to establish the possible genetic divergence between the Sierra de G r e d o s slopes, a n d to clarify the i m p o r t a n c e of diverse factors in the differentiation of these p o p u l a t i o n s within the I b e r i a n variation.


Acknowledgements This study was s u p p o r t e d by a grant from the ' D i r e c c i 6 n G e n e r a l de Investigaci6n Cientifica y T6cnica', PB88-0119. T h a n k s to R o b i n Rycroft for helping with the English manuscript. T h a n k s also to the ' H e r m a n d a d de D o n a n t e s de Sangre Hospital N u e s t r a Sefiora de Sonsoles Avila'.

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Genetic p o l y m o r p h i s m s in Gredos, Central Spain

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MORENO, P., and MORAL, P., 1983, Ditribucidn de grupos sanguineos y enzimas eritrocitarios en una poblacion de Gerona (Espafia). Garcia de Orta Serie Antropolobiologia, 2, 87-92. PANADERO, A. M., 1988, Anfilisis de la variabilidad cualitativa de componentes del complemento en la poblaci6n de Barcelona. PhD thesis, University of Barcelona. PANADERO, A. M., MORENO, P., and MORAL, P., 1986, Variantes gen~ticas de los componentes C3, C6 y BF del sistema complemento en el Pallars Sobir/t. Trabajos de Antropologla, 20, 373 389. PANCORBO, M. M., MAZON, L. I., and LOSTAO,C. M., 1986, A cline in the acid phosphatase-I distribution in the Iberian Peninsula. Annals of Human Biology, 13, 297 300. REGUEIRO, J. R., and ARNAIZ-VILLENA,A., 1984, C3 polymorphism, HLA and chronic renal failure in Spaniards. Human Genetics, 67, 437-440. REYNOLDS, J., WEIR, B. S., and COCKERHAM, C. C., 1983, Estimation of the coancestry coefficient: Basis for a short-term genetic distance. Genetics, 105, 767 779. SANDIUMENGE,T., VIVES,S., and MORAL, P., 1993, Orosomucoid and haptoglobin polymorphisms in two Spanish Pyrenean populations. Gene Geography, 7, 243-250. TEISBERG, P., 1970, High voltage agarose gel electrophoresis in the study of C3 polymorphism. Vox Sanguinis, 19, 47-56. TROT1RO, M. A., 1990, Gredos. La Sierra y su entorno (Madrid de Obras Pflblicas y Urbanismo. Instituto deI Territorio y Urbanismo), Chapter III, pp. 75-149. WORKMAN, P. L., and N1SWANDER, J. D., 1970, Population studies on south-western Indian tribes. II. Local genetic differentiation in the Papago. American Journal of Human Genetics, 22, 24-49. Address for correspondence: Dr Pedro Moral, Laboratorio de Antropologia, Departamento de Biologia Animal, Faculdad de Biologia, Universidad de Barcelona, Av. Diagonal 645, 08028 Barcelona, Spain. Zusammenfassung. An zwei Stichproben autochthoner Bev61kerungen, die beiderseits des Gebirgszuges Sierra de Gredos in Zentralspanien leben, wurden die Variationen in den Serumproteinen Orosomucoid (ORM 1) und 3. Komponente des KompIements (C3) sowie den erythrozyt/iren Enzymen 6-PhosphogluconatDehydrogenase (G6PD), Adenosindesaminase (ADA), Esterase D (ESD), und Saure Phosphatase (ACP1) untersucht. Die Ergebnisse sprechen ffir lediglich eine moderate Difl'erenzierung zwischen den zwei Gredos-Subpopulationen, die im Zusammenhang mit anderen Bev61kerungsgruppen der iberischen Halbinsel diskutiert wird. Ffir die meisten der untersuchten Marker wichen die Allelfrequenzen in Gredos deutlich von denen ffir andere iberische Bev61kerungen ab: relativ hohe C3"S, ORMI*F, ADA*t, ESD*2 und ACPI*2 Frequenzen sind charakteristisch f/ir die Gredos-Stichproben. Diese Differenzierung war ffir die n6rdliche Gredos Population ausgepr/igter, was mit den geographischen Besonderheiten delc Region in Zusammenhang gebracht werden konnte. Resum& Les variations du troisi6me compl6ment humain (C3), des prot6ines du serum orosomucoides (ORM1), des enzymes 6rythrocytaires 6-phosphogluconate d6hydrog6nase (6PGD), ad6nosine deaminase (ADA), est&ase D (ESD) et acide phosphatase (ACP1), ont 6t6 examin6es dans deux 6chantillons de populations autochtones vivant sur chaque versant de la cha~ne montagneuse de la Sierra de Gredos darts le centre de l'Espagne. Ces r6sultats n'indiquent qu'une faible diff6renciation des sous-populations de Gredos, qui est discut6e en relation avec d'autres groupes de la p6ninsule ib6rique. Les fr6quences all~liques de Gredos montrent une hdtdrogdndit6 remarquable par rapport aux autres populations pour la plupart des marqueurs 6tudi6s; des fr~quences g~niques relativement 61ev6es de C3"S, ORMI*F, ADA*I, ESD*2 et ACPI*C sont caract6ristiques des groupes de Gredos. Cette diff&ence est plus marqu6e dans la population de la pattie nord de Gredos et pourrait etre associ6e aux particutaritds g6ographiques de cette r6gion.