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1、2021/3/10授课:XXX1 Apple (Malus domestica Borkh.) a.k.a. Malus pumila P. Mill. Image from Jake Fleming Department of Geography, UW Madison 2021/3/10授课:XXX2 Family: Rosaceae Subfamily:Maloideae (with pears) Genus:Malus (40 sp.) Section:Malus Series:Malus Species:domestica Taxonomy A n
2、ote on nomenclature: In his 2006 book The Story of the Apple, BE Juniper refers to both the domestic apple and the wild Central Asian apple as Malus pumila. The USDA has also adopted this convention. For clarity, and as Coart et al. 2006 calls into question the hypothesis upon which this is based, I
3、 use the older names: Malus sieversii for the wild Central Asian apple and Malus domestica for the orchard apple. 2021/3/10授课:XXX3 Importance Worlds most important temperate fruit crop: 63 million tons/yr. Leading producers in 2004: China 18.7 Mtons (4 Mtons in 1990), United States 6 Mtons, Russia,
4、Germany, Japan Food and Agriculture Organization of the United Nations. 2021/3/10授课:XXX4 Malus domestica - the specifics Woody, long-lived tree Unlike congeners, extreme heterozygosity, does not breed true. Single parental event yields massive variation in fruit color, size, ta
5、ste, flower color, thorns, tree habit, so Cultivars must be vegetatively propagated “instant domestication” Perfect, self-incompatible flowers. n = 17. Most congeners and cultivars 2n, some 3n, 4n. 2021/3/10授课:XXX5 3 Stories of Domestication Compilospecies conventional wisdom until c. 1990. Eurasian
6、 origin, somewhere/everywhere Malus sieversii Roem. Vavilov 1930. Harris, Robinson, Juniper 2002. Central Asian origin BREAKING! Malus sylvestris Mill. Coart et al. 2006. European origin? 2021/3/10授课:XXX6 Compilospecies hypothesis Hypothesis: Malus domestica arose from some combination of crabapples
7、 in Eurasia, with possible recent introgression in North America (Watkins 1995) In Maloideae, hybridization between genera not rare (e.g. Malus x Pyrus) In Malus, species boundaries fuzzy, some hybrids occur M. domestica may (or may not) readily hybridize with sympatric congeners 2021/3/10授课:XXX7 M.
8、 sieversii hypothesis As the Tien Shan Mountains rose and the Gobi and Taklamakan Deserts grew, ancestral Malus populations were isolated Among a very diverse population making up as much as 80% of the forest, Some wild fruits are indistinguishable from cultivars Selection by bears? Human-mediated d
9、ispersal along Silk Road trade routes (but not by current residents) (Juniper and Mabberley 2006) Hypothesis: Malus sieversii of Central Asia is the wild ancestor of domesticated apples. Other species contributed little or nothing. (Vavilov 1930) 2021/3/10授课:XXX8 M. sieversii hypothesis As the Tien
10、Shan Mountains rose and the Gobi and Taklamakan Deserts grew, ancestral Malus populations were isolated Among a very diverse population making up as much as 80% of the forest, Some wild fruits are indistinguishable from cultivars Selection by bears? Human-mediated dispersal along Silk Road trade rou
11、tes (but not by current residents) (Juniper and Mabberley 2006) Hypothesis: Malus sieversii of Central Asia is the wild ancestor of domesticated apples. Other species contributed little or nothing. (Vavilov 1930) 2021/3/10授课:XXX9 2021/3/10授课:XXX10 M. sieversii hypothesis As the Tien Shan Mountains r
12、ose and the Gobi and Taklamakan Deserts grew, ancestral Malus populations were isolated Among a very diverse population making up as much as 80% of the forest, Some wild fruits are indistinguishable from cultivars Selection by bears? Human-mediated dispersal along Silk Road trade routes (but not by
13、current residents) (Juniper and Mabberley 2006) Hypothesis: Malus sieversii of Central Asia is the wild ancestor of domesticated apples. Other species contributed little or nothing. (Vavilov 1930) 2021/3/10授课:XXX11 2021/3/10授课:XXX12 2021/3/10授课:XXX13 M. sieversii hypothesis As the Tien Shan Mountain
14、s rose and the Gobi and Taklamakan Deserts grew, ancestral Malus populations were isolated Among a very diverse population making up as much as 80% of the forest, Some wild fruits are indistinguishable from cultivars Selection by bears? Human-mediated dispersal along Silk Road trade routes (but not
15、by current residents) (Juniper and Mabberley 2006) Hypothesis: Malus sieversii of Central Asia is the wild ancestor of domesticated apples. Other species contributed little or nothing. (Vavilov 1930) Bearington Bears catalog. $10.95 2021/3/10授课:XXX14 M. sieversii hypothesis As the Tien Shan Mountain
16、s rose and the Gobi and Taklamakan Deserts grew, ancestral Malus populations were isolated Among a very diverse population making up as much as 80% of the forest (adaptive?), Some wild fruits are indistinguishable from cultivars Selection by bears? Human-mediated dispersal along Silk Road trade rout
17、es (but not by current residents) (Juniper and Mabberley 2006) Hypothesis: Malus sieversii of Central Asia is the wild ancestor of domesticated apples. Other species contributed little or nothing. (Vavilov 1930) Bearington Bears catalog. $10.95 2021/3/10授课:XXX15 Geography of M. sieversii Harris et a
18、l. 2002 Note: Whats up with this map? 2021/3/10授课:XXX16 Dispersal into Europe Silk Road was operational by 2100 ybp. In summer, went straight through Tien Shan One big apple found in Ireland, 3000 ybp. What is it? Alexander the Great, 2300 ybp, fought mock battles with apple projectiles. Must have b
19、een bigger than crabs. 2400 ybp, Celt-Persian contact? 1300 ybp, Muslim empire, Central Asia-Spain 2021/3/10授课:XXX17 Molecular evidence Harris et al 2002 Wild apple: Malus sieversii Domesticated apple: Malus domestica cDNA gene matK Only 16 of 1341 characters informative 18-bp duplication in M. dome
20、stica and 1 M. sieversii accession Other M. sieversii accessions tested only for duplication, did not have it 2021/3/10授课:XXX18 Nuclear ribosomal ITS Robinson et al 2001 No M. sylvestris = 2021/3/10授课:XXX19 USDA collections, M. sieversii Forsline et al. 2003 2021/3/10授课:XXX20 Harris et al 2002s matK
21、 duplication Forsline et al. 2003 8 of 10 M. sieversii accessions from Uzbekistan and Tajikistan (Robinson et al 2001) 2021/3/10授课:XXX21 Other genetic support for M. sieversii Morphological + RAPD + sequence data (ITS1, 5.8S rRNA, ITS2, matK) Forte et al. 2002 Isozymes DIA-2, AAT-2, PGM-1 and PGM-5
22、Wagner and Weeden 1999 Sampling? Strength of support? I dont know Forte, A.V. et al. 2002. Phylogeny of the Malus (apple tree) species, inferred from the morphological traits and molecular DNA analysis. Russian Journal of Genetics 38: 1150-1160. Wagner, I. and Weeden, N.F. 2000. ISOZYMES IN MALUS SY
23、LVESTRIS, MALUS DOMESTICA AND IN RELATED MALUS SPECIES. Acta Hort. (ISHS) 538:51-56 2021/3/10授课:XXX22 Floral morphology Fruit morphology Morphological evidence for M. sieversii Juniper and Mabberley 2006 2021/3/10授课:XXX23 Hybridization b/w cultivars and sympatric European crab apple, M. sylvestris,
24、“almost absent” Coart et al. 2003 Molecular evidence against M. sylvestris 3 of 76 “wild” specimens STRUCTURE 2021/3/10授课:XXX24 BUT WAIT. M. sylvestris Hypothesis: Actually, M. sylvestris, the European crab apple, is in some way ancestral to domesticated apples Unlike M. sieversii, M. sylvestris is
25、solitary, not notably heterogeneous, rare, and produces bitter, inedible fruit. Contrary to their own earlier work, Coart et al.s JULY 2006 article shows M. sylvestris to be much closer to M. domestica than previous, using cDNA PCR-RFLP 16 different chloroplast haplotypes based on: matK duplication,
26、 1 point mutation, 2 restriction endonucleases (EcoRI, MseI) 2021/3/10授课:XXX25 Coart et al. 2006 2021/3/10授课:XXX26 Coart et al. 2006 2021/3/10授课:XXX27 Coart et al. 2006 2021/3/10授课:XXX28 Coart et al. 2006 matK dupII 2021/3/10授课:XXX29 Greater chloroplast diversity of cultivars suggests some kind of h
27、ybridization Geographic distribution of rare haplotypes suggests sylvestris x domestica hybridization Also, Coart et al 2006 2021/3/10授课:XXX30 Coart et al. 2006 2021/3/10授课:XXX31 Selected Sources Coart, E.L.S., et al. 2003. Genetic variation in the endangered wild apple (Malus sylvestris (L.) Mill.) in Belgium as revealed by amplified fragment length polymorphism and microsatellite markers. Molecular Ecology 12: 845-857. Coart, E.L.S., et al. 2006. Chloroplast diversity in th
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