Báo cáo khoa học: "Genetic variation in European larch (Larix decidua Mill)"

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Original article Genetic variation in European larch (Larix decidua Mill) J Maier Universität München, Lehrstuhl für Forstbotanik, Amalienstraße 52, 8000 Munich 40, Germany (Received 22 February 1991; accepted 25 September 1991) Summary — Levels of electrophoretically demonstrable diversity of 7 allozyme loci were estimated in 7 populations representing the natural range of Larix decidua (Mill). On average the gene diversity was 0.223 and the number of alleles per locus was 2.28. Only 5.1% of the total genetic diversity re- sided among populations with a mean genetic distance among populations of 0.029. The popula- tions could be assigned to two geographic groups, a large one containing populations from the east- ern Alps as well as from Poland and Czechoslovakia and a rather restricted one with a single population from the Western Alps. The most homogeneous populations are those from eastern Eu- rope (Poland, Sudetan region and Tatra mountains). Larix decidua / allozymes / genetic variation Résumé — Variation génétique du mélèze d’Europe (Larix decidua Mill). Le niveau de diversité de 7 marqueurs génétiques a été étudié en électrophorèse pour 7 populations représentatives de la variation naturelle du Larix decidua (Mill). En moyenne, la diversité génétique était de 0,223 et le nombre d’allèles par marqueur 2,28. Seulement 5,1% de la diversité génétique totale étaient repré- sentés dans des populations d’une distance génétique moyenne entre populations de 0,029. Les po- pulations purent être assignées à 2 groupes géographiques, dont l’un, le plus important, comprend les populations des Alpes orientales, ainsi que celles de Pologne et de Tchécoslovaquie, alors que l’autre, plus réduit, ne comprend qu’une seule population des Alpes occidentales. Les populations les plus homogènes sont celles d’Europe orientale (Pologne, Sudètes, chaîne du Tatra). Larix decidua / diversité génétique /marqueurs génétiques
INTRODUCTION liak et al, 1988) and for L occidentalis (Fins and Seeb, 1986). However, for L decidua from Poland segregation studies of allo- Larix decidua Mill has its natural range zyme loci are already available (Mejnarto- restricted to four distinct areas of Central wicz and Bergmann, 1975; Kosinski and and eastern Europe: the Alps, the Sudetan Szmidt, 1984; Lewandowski and Mejnarto- region, the Tatra Mountains and scattered wicz, 1990a, b, 1991).In this study seeds throughout Poland (fig 1).Ithas been the from provenances covering all four parts of subject of both numerous provenance ex- the range of European larch were analy- periments (Cieslar, 1899, 1914; Varma, sed by gel electrophoresis. The genetic in- 1949; Leibundgut, 1959; Barnes, 1977; terpretation of six enzyme gene markers Giertych, 1979; Schober, 1977, 1985) and as well as the calculation of genetic differ- other studies concerning phenotypic traits entiation parameters will be presented. (Bouvarel and Lemoine, 1958; Gathy, 1959; Schreiber, 1960, 1961, 1963; Kral, 1966, 1967; Simak, 1967; Lang, 1976; Lei- MATERIALS AND METHODS bundgut, 1985). These investigations un- derline the fact that European larch is a Geographic location and background informa- highly variable species. Allozyme surveys tion for the seven populations of Larix decidua estimating gene diversity among popula- are given in figure 1 and table I. Bulked seed tions are not known for Larix decidua, but samples, where seeds from several trees were pooled, as well as seed lots from individual trees within the genus Larix for L laricinia (Che-
sampled. Both of them originate from indi- number 1 being assigned to the most mobile were genous populations; the former were provided band at any locus. A locus was considered poly- by forest research stations in Poland, Czechos- morphic if more than one allele was observed, lovakia, and Austria, and the latter were collec- regardless of allelic frequencies. ted by us. Seeds from many small stands from Nei’s (1972) genetic distance (D) was used the Sudetan region as well as from a limited quantify the degree of differentiation among to area in the eastern Alps were grouped, respecti- populations. Cluster analysis, using the vely. UPGMA-method, was performed with SPSS Electrophoretic analysis was carried out on (Norusis, 1986) on the matrix of Nei’s genetic the endosperm. Between 50 and 144 have been distances. Gene diversity analysis was calcula- examined for the bulk provenance collections. ted according to Nei (1973). A measure of total For the provenances with single tree seeds, six gene diversity is H 1 - Σp where p is the 2 , i 2 T = endosperms per tree were analysed. Horizontal frequency of the i th of k alleles. H par- T is mean starch gel electrophoresis was carried out to se- titioned in H H + D where H D TSST =, S ST and are parate isozymes for six enzyme systems: average gene diversities within and among po- pulations, respectively. G the proportion of ST is interpopulation gene diversity H . T RESULTS IDH and GDH found to be monomor- were Details of gel and electrophoresis buffer and phic. No variation was observed for the al- staining mixtures were taken from Conkle et al lozyme encoded by these loci (fig 2a, b). (1982) and from Müller-Starck (personal com- munication). Enzyme band phenotypes, evi- Two zones of activity were observed on dence from gametic segregation ratios and gels stained for G6PDH. The lower zone close analogy to results from other isozyme in- stained inconsistently and was therefore vestigations in larch (Cheliak and Pitel, 1985; not scored. The fastest migration zone ex- Fins and Seeb, 1986; Lewandowski and Mejnar- hibits 3 bands differing in mobility (fig 2c). towicz, 1990a, b; Ying and Morgenstern, 1990) Thus, 1 locus with 3 alleles was postula- were the basis for genetic interpretation of the ted. A heavily stained zone with 2 single- zymograms. Capital letters refer to gene loci,
for SKDH showed 2 zones of activity. The faster zone was unreliably stained and could not be considered. The more catho- dal zone exhibits 5 bands differing in mobi- lity and staining intensity. In addition, a null allele was observed. Thus, this zone was interpreted as 1 locus with 6 alleles (fig 2e). A 5 zone banding pattern, inferred as four loci and an interlocus heterodimer, was recorded for MDH. On account of poor band resolution, MDH-B and MDH-C were not further analysed. Both MDH-A and MDH-D performed 3 bands suggesting 2 loci with 3 alleles each (fig 2f and 2g). Allele frequencies are given in table II. banded and 1 double-banded phenotype When comparing gene frequencies of the found for 6PGDH, suggesting a 3- 7 populations, qualitative differences occur was only in rare variants. The same allele pre- allele locus (fig 2d). Gametophytes scored
vails at every locus in all populations with the exception of the G6PDH. The frequen- cy of the G6PDH-1 allele exceeds that of G6PDH-2 at the provenances Pfitsch and Vintschgau, while for the rest the opposite is true. The estimates of genetic distances for all combinations of provenances averaged over the 7 loci are presented in table III. The distances (average 0.029) are lowest plon population to 0.260 in the Ostalpen among the eastern European samples population with a mean of 0.223 over all from Poland, the Sudetan region and the (E-Alps) populations. The mean number of Tatra Mts (0.004-0.009). The Simplon po- alleles per locus was 2.28 with a minimum pulation appeared to be the most divergent of 2.0 in the Pfitsch and Simplon popula- from all other populations with genetic dis- tions and a maximum of 2.6 in the Sudetan tances rising up to 0.099. All alpine region. On the average, 94.9% (Hs/Hr x samples have relatively large average dis- 100%) of the gene diversity resided within tances between each other. stands and 5.1% among stands (G ST x Figure 3 shows the dendrogram resul- 100%, table V). ting from UPGMA clustering based on NEI’s genetic distance. The general analy- sis showed that two large groups were de- DISCUSSION lineated. The Simplon material from the western Alps is clearly distinct from the other 6 populations. In this cluster contai- Monomorphic loci at GDH are reported for ning 6 populations the 3 provenances from L laricina (Cheliak and Pitel, 1985; Ying eastern Europe form a relatively homoge- and Morgenstern, 1990) and at IDH for neous sub-cluster which is slightly different L laricina (Cheliak and Pitel, 1985) as well from the Alpine provenances. as for L occidentalis (Fins and Seeb, Gene diversity per locus varies widely 1986). Lewandowski and Mejnartowicz from 0.027 at MDH-A to 0.653 at G6PDH (1990a) found these 2 enzyme systems (table IV). The mean heterozygosity per controlled by 1 locus with 1 dominating al- population ranges from 0.140 in the Sim- lele and 2 rare alleles each. Corresponding
1983; Cheliak and Pitel, 1985; Yeh et al, to the present results a 3-allele locus at 1985; Fins and Seeb, 1986; Ernst et al, G6PDH was found for L decidua (Lewan- dowski and Mejnartowicz, 1990a) and for 1987; Merkle and Adams, 1987; Berg- L laricina (Cheliak and Pitel, 1985). At mann, 1988; El-Kassaby, 1989; Lewan- dowski and Mejnartowicz, 1990a) are in SKDH Lewandowski and Mejnartowicz agreement with the present results. For L (1990a) detected 1 locus with four alleles. decidua Lewandowski and Mejnartowicz Further studies of SKDH in Larix have (1990a) observed at MDH1 (MDH-A) and been done on one single clone of both Eu- MDH4 (MDH-D) a deviating number of al- ropean and Japanese larch (Bergmann and Ruetz, 1987). The finding of four MDH leles. For Polish larch however, the postu- loci in many other conifers (Wheeler et al, lation of a monomorphic locus at MDH-D
II: Population Lysa Gora) is consis- larch populations. The isolated position of (table tent with observations by Lewandowski the Simplon stand in genetic distance ma- Mejnartowicz (1990b). This indicates trix and in cluster analysis may be due to and that deviation in number of alleles per the relatively small sample size (23 trees). locus at GDH, IDH, MDH and SKDH in pre- However, cluster analysis derived from sent results compared to those by Lewan- monoterpene data of larch seedlings (un- dowski and Mejnartowicz (1990a) may be published data) exhibits full correspon- dence to the isozyme results. In addition, due to different populations investigated. At 6PGDH one (Cheliak and Pitel, 1985) or provenance experiments support the re- sults from both the isoenzyme and resin oil two (Fins and Seeb, 1986) polymorphic loci had been reported for L laricina and L analyses. Substantial differences in growth rate between western and eastern Alpine occidentalis, respectively. provenances have been pointed out (Scho- Levels of genetic distances and gene di- ber, 1977, 1985). Considerable amounts of versity among provenances of L decidua differentiation in several traits among pro- reveal very similar average values for L la- resulted in several authors venances ricina (Cheliak et al, 1988); however, the speaking of alpine larch races and/or eco- average number of alleles per locus is hi- types (Wettstein, 1946; Rubner, 1954; gher in L decidua. On the other hand, Mayer, 1961; Kral, 1967; Leibundgut, L occidentalis (Fins and Seeb, 1986) dif- 1985). In contrast, the eastern European fers considerably from L laricina and L de- provenances form a quite uniform group in cidua by relatively low genetic variability as respect to larch canker susceptibility and well as genetic distances among popula- growth rate (Schober, 1977, 1985). Accor- tions. This is surprising considering the ex- ding to these and present results, the tent of the species ranges. L laricina ex- larches from Poland, the Tatra Mts and the pands continent-wide, while L decidua and Sudetan region may be regarded as one L occidentalis are localized in restricted re- single race. gions. Fins and Seeb (1986) suggest that low genic differentiation and diversity among stands of western larch may be the ACKNOWLEDGMENTS result of isolated refugia during Pleisto- cene glaciation and founder effects after fires. On the other hand, with regard to the indebted to P Schütt and G Aas for review- I am relatively low number of enzyme systems the manuscript. I also wish to thank JM Vin- ing cent, A Stapf and R Schacher for linguistic ad- asseyed, the results for L decidua should vice and R Vogtmann for drawing figures. This be interpreted carefully. Nevertheless, a study was financially supported by a grant from substantial difference in gene diversity bet- the Deutsche Forschungsgemeinschaft, Bad ween L decidua and L occidentalis re- Godesberg. mains. Genetic variability of L decidua evalua- ted for two Polish stands and for a seed or- REFERENCES chard in Poland was considered to be low (Mejnartowicz and Bergmann, 1975; Ko- Barnes BV (1977) The International Larch Pro- sinski and Szmidt, 1984). In this study no venance Test in southeastern Michigan, obvious low gene diversity in the Polish po- USA. Silvae Genet 26, 145-148 pulation (Lysa Gora) was found. Bergmann F (1988) Untersuchungen der gene- Nei’s (1972) genetic distance indicated tischen Differenzierung der Fichte in den Seetaler Alpen an Enzym-Genloci. Forum- relatively large genetic differences among
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