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1、Apple (Malus domestica Borkh.) a.k.a. Malus pumila P. Mill. Image from bestapplesJake FlemingDepartment of Geography, UW MFamily: RosaceaeSubfamily:Maloideae (with pears)Genus:Malus (40 sp.)Section:MalusSeries:MalusSpecies:domesticaTaxonomyA note on nomenclature: In his 2019 b

2、ook 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. 2019 calls into question the hypothesis upon which this is based, I use the older names: Malus siever

3、sii for the wild Central Asian apple and Malus domestica for the orchard apple.billnymanartImportance Worlds most important temperate fruit crop: 63 million tons/yr. Leading producers in 2019: China 18.7 Mtons (4 Mtons in 1990), United States 6 Mtons, Russia, Germany, JapanFood and Agriculture Organ

4、ization of the United Nations. 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, taste, flower color, thorns, tree habit, so Cultivars must be veget

5、atively propagated “instant domestication” Perfect, self-incompatible flowers. n = 17. Most congeners and cultivars 2n, some 3n, 4n. 3 Stories of Domestication Compilospecies conventional wisdom until c. 1990. Eurasian origin, somewhere/everywhere Malus sieversii Roem. Vavilov 1930. Harris, Robinson

6、, Juniper 2019. Central Asian origin BREAKING! Malus sylvestris Mill. Coart et al. 2019. European origin? Compilospecies hypothesisHypothesis: Malus domestica arose from some combination of crabapples in Eurasia, with possible recent introgression in North America (Watkins 2019)In Maloideae, hybridi

7、zation between genera not rare (e.g. Malus x Pyrus)In Malus, species boundaries fuzzy, some hybrids occurM. domestica may (or may not) readily hybridize with sympatric congenersM. sieversii hypothesis As the Tien Shan Mountains rose and the Gobi and Taklamakan Deserts grew, ancestral Malus populatio

8、ns 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 current residents) (Juniper and Mabberley 2019)Hypothesis: Malus siever

9、sii of Central Asia is the wild ancestor of domesticated apples. Other species contributed little or nothing. (Vavilov 1930) M. 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 mak

10、ing 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 current residents) (Juniper and Mabberley 2019)Hypothesis: Malus sieversii of Central Asia is the wild ancestor of domestic

11、ated apples. Other species contributed little or nothing. (Vavilov 1930) M. 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 fruit

12、s are indistinguishable from cultivars Selection by bears? Human-mediated dispersal along Silk Road trade routes (but not by current residents) (Juniper and Mabberley 2019)Hypothesis: Malus sieversii of Central Asia is the wild ancestor of domesticated apples. Other species contributed little or not

13、hing. (Vavilov 1930) M. 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

14、bears? Human-mediated dispersal along Silk Road trade routes (but not by current residents) (Juniper and Mabberley 2019)Hypothesis: Malus sieversii of Central Asia is the wild ancestor of domesticated apples. Other species contributed little or nothing. (Vavilov 1930) Bearington Bears catalog. $10.9

15、5M. 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 (adaptive?), Some wild fruits are indistinguishable from cultivars Selection by bears? H

16、uman-mediated dispersal along Silk Road trade routes (but not by current residents) (Juniper and Mabberley 2019)Hypothesis: Malus sieversii of Central Asia is the wild ancestor of domesticated apples. Other species contributed little or nothing. (Vavilov 1930) Bearington Bears catalog. $10.95Geograp

17、hy of M. sieversiiHarris et al. 2019Note: Whats up with this map?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. Mu

18、st have been bigger than crabs. 2400 ybp, Celt-Persian contact? 1300 ybp, Muslim empire, Central Asia-SpainMolecular evidenceHarris et al 2019Wild apple: Malus sieversiiDomesticated apple: Malus domestica cDNA gene matK Only 16 of 1341 characters informative 18-bp duplication in M. domestica and 1 M

19、. sieversii accession Other M. sieversii accessions tested only for duplication, did not have itNuclear ribosomal ITSRobinson et al 2019 No M. sylvestris=USDA collections, M. sieversiiForsline et al. 2019Harris et al 2019s matK duplicationForsline et al. 20198 of 10 M. sieversii accessions from Uzbe

20、kistan and Tajikistan(Robinson et al 2019)Other genetic support for M. sieversii Morphological + RAPD + sequence data (ITS1, 5.8S rRNA, ITS2, matK) Forte et al. 2019 Isozymes DIA-2, AAT-2, PGM-1 and PGM-5 Wagner and Weeden 2019 Sampling? Strength of support? I dont know Forte, A.V. et al. 2019. Phyl

21、ogeny 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 SYLVESTRIS, MALUS DOMESTICA AND IN RELATED MALUS SPECIES. Acta Hort. (ISHS) 538:51-56 Floral morp

22、hology Fruit morphologyMorphological evidence for M. sieversiiJuniper and Mabberley 2019Hybridization b/w cultivars and sympatric European crab apple, M. sylvestris, “almost absent”Coart et al. 2019Molecular evidence against M. sylvestris3 of 76 “wild”specimensSTRUCTUREBUT WAIT. M. sylvestrisHypothe

23、sis: Actually, M. sylvestris, the European crab apple, is in some way ancestral to domesticated applesUnlike M. sieversii, M. sylvestris is solitary, not notably heterogeneous, rare, and produces bitter, inedible fruit.Contrary to their own earlier work, Coart et al.s JULY 2019 article shows M. sylv

24、estris to be much closer to M. domestica than previous, using cDNA PCR-RFLP 16 different chloroplast haplotypes based on: matK duplication, 1 point mutation, 2 restriction endonucleases (EcoRI, MseI)wikipediaCoart et al. 2019Coart et al. 2019Coart et al. 2019Coart et al. 2019matK dupII Greater chlor

25、oplast diversity of cultivars suggests some kind of hybridization Geographic distribution of rare haplotypes suggests sylvestris x domestica hybridizationAlso, Coart et al 2019Coart et al. 2019Selected SourcesCoart, E.L.S., et al. 2019. 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. 2019. Chloroplast diversity in the genus Malus: new insights into the re

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