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In situ and ex situ assessment of morphological and fruit variation in Scandinavian sweet cherry
Inger Hjalmarssona, Rodomiro Ortizb,*
È gen 2, PO Box 41, S-230 53 Alnarp, Sweden The Nordic Gene Bank, Smedjeva Department of Agricultural Sciences, The Royal Veterinary and Agricultural University, 40 Thorvaldsensvej, DK-1871 Frederiksberg C, Denmark Accepted 17 September 1999
Abstract Sweet cherry is a tall, deciduous tree producing stone fruits. This diploid outcrossing species was domesticated in Asia but has been grown in orchards and home gardens in Scandinavia for many years. In situ and ex situ assessments of phenotypic variation in sweet cherry accessions were performed to determine the reliability of such assessments, and to determine relationships between Nordic populations. Principal component analysis (PCA) based on in situ data revealed that accessions were mostly clustered according to their country of origin. PCA based on ex situ assessment of accessions that were propagated by seed at Hornun (Denmark) did not agree with the PCA based on in situ data. These contrasting results suggest that phenotypic assessment in sweet cherry depends on the environment, genotype, and the interaction between them. Phenotypic diversity accounted for by in situ assessment may not be always true, while phenotypic differences determined by ex situ assessment may be confounded by the genotype-by-environment interaction, or could depend on the new genotypes arising from open pollination after seed propagation. Our research also suggests that ecotype differentiation could occur in wild Scandinavian sweet cherry. Fruit descriptors were among the best to distinguish between Scandinavian populations. Previously reported monogenic characteristics showed intermediate narrow-sense heritability, as suggested by the percentage of total variation accounted by the half-sib populations. # 2000 Elsevier Science B.V. All rights reserved. Keywords: Prunus avium; Conservation genetic resources; Phenotypic diversity
Corresponding author. Present address: International Crops Research Institute for the Semi-Arid Tropics (ICRISAT), Patancheru 502 324, Andhra Pradesh, India. Tel.: 91-40-3296161; fax: 9140-3296182. E-mail address: email@example.com (R. Ortiz) 0304-4238/00/$ ± see front matter # 2000 Elsevier Science B.V. All rights reserved. PII: S 0 3 0 4 - 4 2 3 8 ( 9 9 ) 0 0 1 2 3 - 5
) provided information for eight distinct cultivars. Ortiz / Scientia Horticulturae 85 (2000) 37±49 1. 1888). to determine the reliability of such assessments. 1996). R. This paper provides analyses of data collected from 36 populations of wild mother trees in Sweden.C. Sweet cherry regenerates well in natural habitats. e. but nowadays trees of this species are observed growing wild in most of Europe and nearby Asia. Probably the ®rst domestication occurred in this region by moving promising wild individuals into gardens. which has not yet been well explored and exploited. and from data recorded in a second open-pollinated generation from some of these mother trees. 1990). Research to obtain more knowledge about the geographic distribution and the biological variation in the Nordic gene pool of wild sweet cherry was performed by the Nordic Gene Bank (Fernqvist. 1996). Iezzoni et al. sweet cherry cultivation seems to be very old.38 I. exhibit high genetic variability. Norway and Bornholm (southern Denmark). According to Bargioni (1996) sweet cherry.D. and to determine the relationships between Nordic populations. So far the ECP/GR has focused mainly on cultivar description.g. It was recommended that wild forms. Sweet cherry cultivation started in Scandinavia years ago (Shu during the Middle Ages. 1988a).. becomes rare towards the north because of its sensitivity to cold winters (Dahl. now IPGRI) started collaborative European genebank research on Prunus at the beginning of the 1980s (Schmidt. An update of this work has been provided by Gass et al. which may be seed propagated but is self-incompatible. 1985). Some of the cherry cultivars grown today are either old local cultivars or their next generation. . is mostly found in single tree stands or in small groups. Findings in Norwegian graves revealed that cherries were part of the Scandinavian diet 1500 È beler. while Pliny (23±79 A. The aims of our paper were to carry out analyses on in situ data and ex situ assessment of the accessions available in the Nordic Gene Bank. (1990) also indicated that knowledge about the inheritance of horticulturally important characteristics is limited. which is vigorous and can be 20 m tall. The Roman writer Varro (116±27 B. recently. The tree. particularly in northern Iran and Ukraine (Webster. The species. landraces and old cultivars should be described. (1996). Hence there is a need for a systematic assessment of cultivars and wild germplasm for speci®c characteristics. as compared to other fruit species. in eastern Europe and the former USSR (Iezzoni et al. Introduction Sweet cherry (Prunus avium L.) was domesticated in Central Asia. Both young seedlings (raised from cross-pollination) and suckers (genetically identical with the mother tree) can be easily transplanted.) described its grafting technique. Hjalmarsson. According to Dahl (1988b). The European Cooperative Programme for Conservation and Exchange of Crop Genetic Resources (ECP/GR) and the International Board for Plant Genetic Resources (IPBGR.
I. In situ assessment Based on Nordic ¯oras (Lagerberg. 2 pink. The environment where these trees were growing was an undulating pasture. 3 medium. 5 very sweet 1 whitish. The Danish and Swedish accessions used were described in 1982 or 1985. and Va È lla (55814H ) and were described at 30 locations. and was therefore chosen for the inventory. There were 19 trees per population. 4 purple mm . 1939.1. while the growing sites in Grimstad were characterized by forests and slopes. In Norway sweet cherry is found in the southern lowland areas. Hjalmarsson. while the Norwegian accessions were described in 1993. È beler. 5 purple. Lagerberg and Holmboe. Among the Danish islands. 3 red on pale yellow ground. The Danish accessions were isolated on the island of Bornholm (50 km away from the nearest mainland). 3 red. areas with wild sweet cherry Shu trees were identi®ed and listed. 2 orange-yellow. Bornholm (55815H ) is well known for sweet cherry growing. 4 sweet. R. In Sweden the species spreads from Scania È rmland (north). 1888) and local horticultural expertise. 2 spreading. 1947. Small populations (1±3 trees) (south) to Uppland. 6 black 1 very acid. 3 tree with more than one stem 1 erect. The shortest distance between the locations of Norwegian and Swedish accessions was approximately 300 km. the southernmost being Baskemo the northenmost at Tullgarn (59818H ). 4 heavy 1 yellow. 3 intermediate. At the time of fruit ripening (July±August) the mother trees were characterized with descriptors recommended for in situ morphological variation (Table 1). The trees were mostly found on ¯at and cultivated land. Table 1 Descriptors considered for the in situ morphological variation between 36 Scandinavian populations of sweet cherry Descriptor Shrub/tree height Habitus 1 Habitus 2 Fruit size Fruit set Fruit colour Fruit taste Fruit flesh colour Stone size Scale m 1 tree. 2 weak. specially near the Oslo ®ord and along the coast. one near Jelùy in the Oslo district (59810H ) and the other near Grimstad (5888H ). 4 red. Mother trees in small populations (1±4 trees) were described at four different locations. In our analysis we used mother trees from two different populations. 2 shrub. Dalarna. 3 drooping mm 1 none. In Jelùy the trees were mostly found near ®eld borders. 2 acid. Ortiz / Scientia Horticulturae 85 (2000) 37±49 39 2. Materials and methods 2.
Hjalmarsson. growth. R. 4 red. The seeds germinated well in autumn 1987. Ex situ assessment Seeds collected from open-pollinated mother trees in 1985 were sown in spring 1986. while growth was the only character not recorded in 1997. 1989). viz. In 1995. and 20 mother trees grown at seven locations in southern Sweden.2. 6 black 1 early. After eight growing seasons berries were observed on most trees. 2. Ortiz / Scientia Horticulturae 85 (2000) 37±49 2.3. 9 susceptible 1 none. canker susceptibility owing to Pseudomonas spp. May 1997 and August 1997. Fruit and stone weight were recorded in sets of 50 fruits. The planting distance within the rows was approximately 1 m and between rows 5 m. 4 elongate g mm g . 3 round. There were 300 Danish derived seedlings (20±60 half-sibs per mother tree) and 317 Swedish derived seedlings (4±20 half-sibs per mother tree). 9 vigorous Count 1 resistant. 9 best Percentage in flower development between green clusters and full bloom Percentage (100 ± percentage white buds) 1 low. 3 late 2 flat round. three of these descriptors. The descriptors used for this ex situ morphological characterization are given in Table 2. the resultant variation can be Table 2 Descriptors considered for the ex situ morphological variation between 28 half-sib populations from Denmark and Sweden Descriptor Growth Stem number Canker Intensity of bloom White buds Full bloom Fruit set Fruit colour Maturity Fruit shape Fruit weight Fruit peduncle Weight 100 stones Scale 1 weak. The half-sib populations were thus characterized in August 1995. The seedlings derived from open-pollinated seed collected from eight mother trees grown at three locations in Bornholm.. If offsprings from each mother tree are progeny tested. and 1680 seedlings were planted in eight rows in an experimental ®eld at Hornum (Denmark). and fruit colour. 9 best 1 yellow. Statistical analysis Analyses of variance (ANOVA) and principal component analyses were carried out in all data sets with the aid of MSTAT-C (Anonymous. were recorded. while length of stem peduncle was measured on 20 samples.40 I. 2 medium.
Thus. size and colour. and these characteristics are those with the highest ei. Likewise. while height. Sweden.05) among the three Scandinavian states for height. the greater the expected gain from selection. First and second principal components (PRIN 1 and PRIN 2. fruit size. habitus 2.I. Each combination consists of a set of weightings known as principal components (PRINs). Principal component analysis (PCA) is a method which can be used to identify patterns in a set of biological data derived from recording several characteristics at a time (Iezzoni and Pritts. The tallest and most erect trees were those of Norway. It seems that fruits become smaller in the north than in the south of Sweden (I. Fernqvist. commun. This half-sib analysis provides indirect information about the amount of additive genetic variation (VA) in the reference material because the covariance between half-sibs is equal to 1/4 VA. which are functions of the eigenvalues (li) and eigenvectors (ei) of the variance/covariance matrix of the original data. The latent roots determine the number of the most important characteristics loading the PRINs. Heritability has been used by plant breeders to calculate gains from selection. 1991).. habitus 1. the characteristics recorded in Bornholm were similar between populations. 3. where VP is the total phenotypic variation. Swedish cherries were larger and more tasty than those from the neighbouring countries. fruit taste and ¯esh colour (Table 3).). the percentage accounted for by the sum of squares owing to the half-sib population was calculated in the ex situ data. The analysis transforms the original correlated measurements into uncorrelated linear combinations of these variables (Hill et al. A descriptor by accession matrix was generated for this PCA and latent vectors were derived from the correlation matrix. pers. SLU. Hjalmarsson. there were signi®cant differences (P < 0. Results 3.. information regarding narrow-sense heritability (h2) of a characteristic may be obtained because h2 VA/VP. colour and taste varied signi®cantly among the two Norwegian populations. fruit set. . Therefore. In situ assessment On average. the higher the heritability. 1998). Successive components (PRIN 3 onwards) accounted for a decreasing proportion of the total variation and were not included in the graphs. and stone size were signi®cantly different among Swedish populations. Ortiz / Scientia Horticulturae 85 (2000) 37±49 41 partitioned into within and between maternal groups (Hill et al. 1998). R. Furthermore. The ¯esh colour was darker in Norway than in Sweden.1. based on the respective list of descriptors. respectively) were plotted to enhance the dispersion of the accessions. fruit set. PCA explains the variance/covariance structure of the data set with a few (usually 2) linear combinations of the original variables. height.
335 0. Both Danish and Norwegian populations are placed towards the left on the PRIN 1 axis.482 0.383 0.478 PRIN 3 0.0 1.5 13.012 1.016 b a b 1.446 0.001.9 0.028 0.038 0.6 9.038 0. The PCA biplot (Fig.231 17. fourth and ®fth principal components of the in situ data assessment of Scandinavian sweet cherry Descriptor Height Habitus 1 Habitus 2 Fruit set Fruit size Fruit colour Fruit taste Latent roots Percentage variance Cumulative variance PRIN 1 À0.048 0.569 0.301 À0. while one of the two Norwegian populations was mixed with those from Denmark.001 0.315 0.6% of the total variance was unevenly loaded.2 2.012 0.179 À0.466 0.146 À0.002 a b 3.428 0.8 3.444 PRIN 5 À0.2 b Colour Taste 4.891 30.0 1.035 0.431 À0.5 3.114 b 0.6 0.905 78.063 0.687 9.9% of the total variation and was unevenly loaded (Table 4). 1) shows Swedish and Danish populations clustered together.5 4.9 12.0 1.4 4.1 a a a 2.282 0. The most important loading characteristics were fruit taste and fruit size as determined by the latent vectors and latent roots. R.797 0.613 0.411 0.189 b Indicates F-test smaller than 1.090 À0.370 0. The ®rst principal component (PRIN 1) accounted for 30.071 0. The most important loading characters were habitus 1 and fruit set as indicated by its eigenvector and latent root.973 13.062 0.807 88.229 1.9 20.441 À0.5 a Flesh colour Stone size 9. third.251 .42 I.417 0.129 À0.0 9.005 0.4 2.380 0.309 À0.539 PRIN 4 0. and thus characterized by their acid Table 4 Eigenvectors for the ®rst.5 3.442 0. second.321 0.498 2.0 1. Ortiz / Scientia Horticulturae 85 (2000) 37±49 Table 3 Average in situ morphological and fruit variation between 36 Scandinavian populations of sweet cherry (descriptor and scale or unit indicated in Table 1) Location Height Habitus 1 Habitus 2 Fruit Set Denmark Norway Sweden P (F-tests) Between states Within Norway Within Denmark Within Sweden a b Size 10.587 48.7 7.002 0.061 64.223 0.162 30. Hjalmarsson.124 16.359 À0.428 0.891 PRIN 2 0. P (F-test) <0.391 0.152 0.612 0. Similarly the second principal component (PRIN 2).379 0.2 1. which accounted for another 17.012 0.3 a 0.008 a a 3.208 0.9 0.
and their PRIN 1 score was higher than those from Danish and Norwegian populations. 1.I. Their relatively high score on the PRIN 2 axis was explained by a speci®c habitus (one stem tree) and heavy fruit set. In this assessment there was no clear cut distinction between Danish and Swedish populations as in the in situ assessment. and maturity and fruit shape in Sweden. R. Similary PRIN 2. The most important positive loading characteristics of PRIN 1 were growth and intensity of bloom. This result was not surprising because of the widespread distribution of sweet cherry in Sweden (about 600 km from south to north). 2 shows the PCA biplot in which arrows indicate the Danish populations. PRIN 1 explained 39. Ortiz / Scientia Horticulturae 85 (2000) 37±49 43 Fig.8% of the total variation and was unevenly loaded (Table 7).05) variation within each country for most characteristics except canker resistance and maturity in Denmark. fruit taste and small fruit size.2. darker colour. but lower fruit weight and weight of 100 stones than Swedish populations (Tables 5 and 6). The most important characteristics of PRIN 2 were fruit colour (negative) and percentage of white buds (positive). The Swedish populations exhibited the greatest variation. Hjalmarsson. Further analysis of variation for each characteristic within Denmark and Sweden showed signi®cant (P < 0. The other characteristics were statistically similar in open-pollinated derived populations collected either in Denmark or Sweden. Fig. higher host resistance to canker. was unevenly loaded. Ex situ assessment The populations from Denmark had on average signi®cantly (P < 0. . Positions of principal component (PC) scores of different Scandinavian accessions of sweet cherry based on in situ assessment.05) larger growth. 3. while the most negative loading characteristic of PRIN 1 was host resistance to canker. which accounted for 21% of the total variation.
44 I.008 a 6.2 0.588 0.006 0.8 3. fourth and ®fth principal components of the ex situ data assessment of Swedish and Danish sweet cherry Descriptor Growth Fruit colour Canker Intensity of bloom Percentage of bloom Number of stems Fruit set Latent roots Percentage variance Cumulative variance PRIN 1 0.435 1.138 À0. third.4 0.564 0.570 0.468 0.037 0.291 0.4 a a 0.441 0.428 À0.006 0.515 0.296 À0.520 0.022 0.418 2.052 60.003 0. second.1 1.8 a 0.474 21.2 4.135 0.220 À0.175 0. Table 7 Eigenvectors for the ®rst.274 0.035 a a Bloom intensity 6.666 0.358 0.013 0. Table 6 Average ex situ fruit variation between 28 Danish and Swedish half-sib populations of sweet cherry (descriptor and scale or unit indicated in Table 2) Population Maturity Fruit Shape Denmark Sweden P (F-tests) Between states Within Denmark Within Sweden a b Weight 1.4 a Weight of 100 stones 147 182 0.598 0.088 93.7 0.085 15.866 PRIN 3 0.261 .356 5.3 1.121 a 3.5 0.001.189 0.309 À0.012 Peduncle 39.180 0.088 0.814 PRIN 2 À0.220 0.004 P (F-test) <0.787 39.192 1.807 88.2 0.4 4. P(F-test) < 0.472 0. Ortiz / Scientia Horticulturae 85 (2000) 37±49 Table 5 Average ex situ morphological variation between 28 Danish and Swedish populations of sweet cherry (descriptor and scale or unit indicated in Table 2) Population Stem Growth Denmark Sweden P (F-tests) Between states Within Denmark Within Sweden a b Number 2.4 23. Indicates F-test smaller than 1.173 PRIN 5 À0.5 5.3 39. Hjalmarsson.367 PRIN 4 0.9 b a a Fruit colour 5.020 0.1 2.188 a Canker 3.814 39.002 b 2.030 À0.439 0.032 À0.033 0. R.501 76.026 0.041 À0.004 Indicates F-test smaller than 1.001.831 0.826 11.229 0.2 b a a White buds (%) 23.2 b a a Fruit set 2.002 0.021 À0.6 6.0 2.
0 8.0 31.5 24.9 31.1 33.5 28.1 10.0 52.2 35.3 64.0 46. Hjalmarsson.2 18.7 300 8 3 HS from Sweden 9.9 35.7 64.6 24.0 20.9 32.8 28.8 56.5 30. Table 8 Percentage of the total variation of ex situ assessment of Swedish and Danish sweet cherry accounted for by the variation among open-pollinated half-sib (HS) populations Characteristic Growth Fruit coloura 1995 1997 Cankera 1995 1997 Intensity of bloom Percentage of white buds Percentage of full bloom Stem number Fruit set Maturity Fruit shape Fruit weight Fruit peduncle Weight of 100 stones Open-pollinated half-sibs Mother trees of HS Locations where mother trees were grown a HS from Denmark 17.0 8. R.4 317 20 7 Combined over years.4 10. 2. .2 30. Ortiz / Scientia Horticulturae 85 (2000) 37±49 45 Fig.6 28.9 22.9 17.2 19.0 27.7 29.2 16.0 25.I.6 15. Positions of principal component (PC) scores of different Swedish and Danish (indicated with arrows) accessions of sweet cherry based on ex situ assessment.
fruit colour.46 I. or could depend on the new genotypes arising from seed propagation after open-pollination.. while the phenotypic variation ascertained by ex situ assessments may be confounded by the genotype-by-environment interaction. Fruit colour seems to be controlled by one gene. date of data recording) and the genotype-by-environment interaction signi®cantly (P < 0. In the ex situ assessment of this investigation. thereby suggesting an intermediate h2 for this characteristic.e. 1996). 1985a). fruit set. Indeed. another fruit species native to Scandinavia (Erstad. This discrepancy could be explained by the in¯uence of the environment. 4. In addition it has a wide natural distribution.001) affected fruit colour and host resistance to canker. the percentage of SS due to HS populations in the combined ANOVA was lower than those from the individual ANOVA. Kolesnikova (1975) (cited by Iezzoni et al. clustering according to geographical origin was not obtained among the open pollinated half-sibs. Discussion Sweet cherry is a species with great phenological and morphological variation. which explains the con¯icting results from the individual and combined ANOVAs for fruit colour and host resistance to canker. and with ``dark'' colour being dominant over ``light'' colour (Theiler-Hedtrich. Hjalmarsson. phenotypic variation determined by in situ assessments may not be always true.. the genotype-by-environment interaction. thereby ecotypes are expected to develop. the populations whose seeds were chosen for deriving seedlings. and the descriptors used for the in situ or ex situ research. This division was based on differences in winter hardiness and fruit quality. For the two characteristics recorded in 1995 and 1997 (fruit colour and resistance to canker). Both the environment (i.. Ortiz / Scientia Horticulturae 85 (2000) 37±49 The percentage of sum of squares (SS) of the analysis of variance accounted by the variation among the open-pollinated half-sibs (HS) was calculated for each descriptor (Table 8).e. The results suggest an intermediate to high heritability for intensity of bloom. Therefore. heritability calculated in single environments (one date for recording or only one location) may be biased upwards. 1990) listed ®ve ecotypes of sweet cherry in the former USSR. fruit taste and size) and fruit set were important factors for clustering. which agreed with the Kolesnikova's ®ndings because good fruit set can only be achieved if trees are hardy enough. R. The pattern of spatial relationship obtained for Scandinavian populations based on the in situ assessment suggests distinct ecotypes of sweet cherry within Scandinavia. In our investigation fruit quality (i. fruit peduncle and stone weight. The other fruit characteristic which also seems to have an intermediate h2 was fruit . The percentage of total variation for fruit colour accounted for by the half-sib populations was on average 30%. for example ecotype differentiation occurs in red currant (Ribes rubrum). Distinct ecotypes are not unique to sweet cherry.
1997). For example. Fernqvist and Huntrieser (1988) Sto used wild sweet cherry as a model for isoenzyme analysis of fruit cultivars and genotypes. which could affect the ex situ assessment in our study. The inheritance to bacterial canker resistance in seedlings was investigated by Theiler-Hedtrich (1985b). Such a procedure provides a means for sequential observations of the clonally propagated germplasm over a long period. Beaver et al. Time of ripening has been suggested as a good cultivar descriptor because known cultivars of sweet cherry ripen over a seven week period. this characteristic appears to have the highest h2 as determined by the percentage of sum of squares accounted by the variation among half-sibs. 1997). who suggested that resistance to this disease was under polygenic control. Perhaps to obtain an accurate assessment of phenotypic variation in sweet cherry it is necessary to study populations over a long period. . Molecular or biochemical characterization may be an alternative for description of wild and cultivated sweet cherry germplasm (Gerlach and È sser. Skin fruit colour has been considered to be in¯uenced by climatic conditions. Fruit set has been reported to be a recessive characteristic because only 5±20% of seedlings obtained were high yielding even if the parental plants were both very good croppers (Theiler-Hedtrich. Ideally. Stylar ribonucleases have also been studied to determine incompatibility within cultivars and progenies of sweet cherry (Boskcovic and Tobutt. and with short length dominant over long (Brown et al. Italian populations of wild sweet cherry were investigated with seven isoenzyme systems (Ducci and Proietti. thereby supplying reliable data for statistical analysis. clones should be propagated and planted in the same testing ®eld and perhaps near the location where the gene bank curator works. has a complex inheritance and the phenotypes for this characteristic may depend on different factors such as hardiness. Consequently. This researcher reported that fruit size and fruit peduncle may be among the most important descriptors for such an assessment. where these populations were situated close to one another. Hjalmarsson. Boskcovic et al. 1996. while the between population variation was high in areas with sparse distribution. Similarly. In our experiment. Propagation should be either through grafting on the same rootstock cultivar or by micropropagation. Ortiz / Scientia Horticulturae 85 (2000) 37±49 47 peduncle. 1994). 1997.. whose variation among half-sib populations accounted in excess of 30%. 1996).. Juice colour was another characteristic suggested by Vittrup Christensen (1970) as a cultivar descriptor to discriminate between coloured and uncoloured fruits. The intrapopulation variability was highest in the northern part of Italy. our single rating of maturity on a 1±3 scale could not have been precise enough. 1995). Such a polygenic system may be affected by the environment and the genotype-by-environment interaction as suggested by our results. Vittrup Christensen (1970) investigated what sweet cherry descriptors could serve to differentiate between cultivars. The characteristic intensity of bloom. R. which has been reported to be simply inherited.I.. Six enzyme systems were enough to identify distinct Scandinavian genotypes.
Brown. T. Hence. 212±218.. Alnarp. i Sverige Pro 4±20. Report 1986±1987. H. S. 81±104. and ex situ conservation in regional seed orchards. Division of Fruit breeding. Report of the working group on Prunus. The Swedish University of Agricultural Pomologi o Sciences. dwar®ng. Fruit Breeding... Inventering och utva È rhamnoides. K. J. 1997. Sper. K. 1996. J. 1997. management and analysis of agronomic research experiments. Selv.). pp. Bot.D.R. 48±50. Boskcovic.. 1988a. Appl. Euphytica 95. C.L.48 I. A. In addition to ®eld gene banks. Cherries. Patterns of random ampli®ed polymorphic DNAs for sweet cherry Gerlach. 1988. 1996. (Eds. Euphytica 88. 75±82. such as host resistance to canker and cracking.. Bargioni. 201±206.F. Ko È rsba È r. H. 245±250. En Pomologi o È ver Dahl. The Swedish University of Agricultural Sciences.. Genetic variability of wild cherry (Prunus avium) in Italy.. and tree size uniformity. C. 73±112.). Rome.F. 1988b. I.A.. New York. References Anonymous. J. A. The Swedish University of Agricultural Sciences.. 1996. È rsba È rstra È dens utbredning och botanik. Tobutt.. I. Sto (Prunus avium L. 213±255. vol. Hjalmarsson. Ist. self-fertility.. I. (Eds. important characteristics for fruit producers. (Ed. 25.. pp. seed germination. 847±852. Erstad..W.R. Beaver. K. Isozyme diversity in sour. Use of isozyme analyses for the identi®cation of fruit cultivars and genotypes. Sweet cherry scions: characteristics of the principal commercial cultivars. Ko È vade Ko È rsba È rssorter. Wiley. ECP/GR.. R. R. Cherries: Crop Physiology. pp. should be considered by gene bank curators in their assessment of variation among sweet cherry accessions. Inheritance of stylar ribonucleases in cherry progenies. R. 1996.). È rsba È rsodlingens utveckling. 1997. sweet. Tobutt.. A. and reassignment of incompatibility alleles to two incompatibility groups..G.. Proietti. Ko È rsba È r.. En Dahl..). Ann. In: Janick. Michigan State University. Fogle. pp. 90. Iezzoni. IPGRI. Huntrieser. I. Ortiz / Scientia Horticulturae 85 (2000) 37±49 Kleinschmit and Stephan (1998) have suggested in situ conservation of natural stands with a minimum of 30±50 individual trees. breeding objectives and methods. Iezzoni. Ramm. K.K. Genet. and ground cherry.) cultivar identi®cation. Ecotype differentation of Ribes rubrum in Norway. N. East Lansing. 1996). Cambridge. Hippophae Fernqvist. 1996. A thorough assessment of wild sweet cherry germplasm will help to obtain clonal archives comprising most of the variability of horticulturally important characteristics. R.E. Angew. R. Alnarp. 1995. Ducci. Russell.N.F. Cambridge University Press. C. In: Fernqvist. Production and Uses.R. In: Webster. 1989.K.. Ê rd. Balsga Gass. 21±23.W. Tobutt. J. Ko È ver i Sverige Pro È vade Ko È rsba È rssorter.... 1996. Fernqvist. I. . F.G. Zanetto. Euphytica 90. 71. È rdering av vildva È xande Prunus-arter och havtorn. MSTAT-C: a microcomputer program for the design. Boskcovic... Correlation of stylar ribonuclease zymograms with the incompability alleles in sweet cherry. Moore.. È sser. Theor. 221±228. In: Fernqvist. A. G.. pp.. Looney. (Ed. Publications-Nordic Gene Bank 31. cryopreservation has also been recommended as a back-up conservation system of genetic resources of Prunus (Gass et al. I: Tree and Tropical Fruits.
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