« »
______ . . , «_____» ______20 ___ .
А А А
" (Abies sibirica Ledeb.)"
06.04.01 –
06.04.01.06 –
______ , . . . . . , , , ______ . . , , ______ . . ., . . . . . , , ,
2020 А
50 , 7 , 9 , 6 , 58 . , , , , , . , (Abies sibirica). . 10 SSR- , - A. sibirica., - , - , .
2
А ...... 3 ...... 4 1 Abies ...... 6 1.1 ...... 7 1.2 ...... 8 1.3 ...... 9 1.4 ...... 12 2 Abies ...... 14 2.1 Abies ...... 14 2.1.1 Abies, ...... 14 2.1.2 Abies, ...... 16 2.2 AFLP – Abies . 17 2.3 Abies, SSR 19 2.4 Abies, SSR ...... 19 2.5 Abies, SSR ...... 20 2.6 Abies, ISSR- ...... 21 3 ...... 22 3.1 (SSRs) ...... 22 3.2 ...... 24 3.3 ...... 25 3.4 ...... 26 4 ...... 29 4.1 ...... 29 4.2 ...... ! . 3
4.3 ! . ...... 31 ...... 33 ...... 40
4
Abies (Mill) , , . , 56 [33]. , , Picea ( ). [31]. . , , [4]. , (Abies sibirica) . ( 6.9 × 10-4) [50], , , , . : , (Abies sibirica). : 1. , . 2. . 3. .
5
. . . . , . ( . . . . . ) ( . . . . . . . ) .
1 Abies
6
1.1
Abies I . . «Elate», 37 . . . , 18 Abies, . , , . , . 1768 . . , 1871 ., , , . 18 Abies [33]. 1928-1929 ., . Abies, 52 12 , 9 , [33]. . . . [7] , « » 1949 ., . . [23] Abies. . . [7] 50 , 40 , 10 - . 31 Abies, 25 . Abies . . [18], (52 ), Abies, 4 : Bracteata, Elate, Abies Piceaster. Abies : Pseudotorreya Abies. 15 , 14 Abies. , 39 , 19 9 . « » , ,
7
. . , [33]. Abies 56 , , - 28, - 4, - 2, - 2, - 5, - 15 [33].
1.2
− Pinophyta. – Pinopsida (Coniferae). - Pinales, (Pinaceae) – ( ) , : (Abietae), (Abies) ( 1) [11].
1 - [11]
9 , Abies sibirica. [13] : - – (Abies sibirica subsp Ledeb.) – ; - – (Abies sibirica subsp. Semenovii) – .
8
, , - ; - , 20 - Abies ×sibiriconephrolepis [24].
1.3
, , , , , ( 2). , [10]. , . – , - , – - , . [20]. Abies . . , , , , [36]. , , , ( ).
9
, , [15]. . ( 2) [11]. - . , , , . , , [11]
10
2 – Abies sibirica: 1 – , 2 – , 3 – , 4 – , 5 – , 6 – , 7 – [11]
, . , [20]. , , , , , . , , , [6]. (Abies sibirica Ledeb.) ( 3). Abies sibirica , - , . Abies sibirica , , ( , ). . , A. sibirica , [5]. , Abies , - , - . . [14].
11
3 - Abies sibirica [3]
1.4
. , . . [19]. [16]. , , , , [17]. , . . .
12
( , ) 1992 - [9]. , A. sibirica . , - , . , Abies sibirica [33].
13
2 Abies
, , , . , [44]. , , , .
2.1 Abies
. . , , . , , [1]. .
2.1.1 Abies,
Abies .
14
. . [28] 11 : (A. sachalinensis): A =
1.57, P100 = 51.5 %, HE = 9.7%. (A. nephrolepsis): A = 1.38,
P100 = 29.1 %, HE = 8.3%. (A. gracilis Kom): A = 1.4,
P100 = 36.5%, Ho = 10.2%, HE = 13.5%.
A. sibirica 15 . HE 6.5% 9.7%, 8%.
100%- P100 13.3% 20% ( 18.9%), A=1.33. . ., [33] 20 A. sibirica
: A = 1.45, P99 = 35%, HE=
6.42%, Ho = 5% ; A. sibirica : A = 1.25, P99 = 20%, HE =
5.65%, Ho = 5.5%. 20 , 11 , . . . . [32], 5 . . : ( ) ( ), – .
: A = 1.35, Ho = 0.044, HE =
0.049, P99 = 25%.
: A = 1.3, Ho = 0.065, HE = 0.072,
P99 = 20 %. 15
. . [22] - ( / A. nephrolepsis Maxim.) (A. sachalinensis Fr. Schmidt) 22 . (A. nephrolepsis): A = 1. 83, P99 = 59.1%, HE = 18.6%, Ho = 18%. (A. sachalinensis): A = 1.71, P99 = 59.1, HE = 17.7%, Ho = 18.6%.
2.1.2 Abies,
. ., 15 , [28]
. Fis 0.013.
Fst 0.093. , 90.7% Abies sibirica , 9.3% . 15 , D , 0 0.049, 0.008. , D (r=0.4488, P=0.002). 11
A. sachalinensis A. nephrolepis (Fst = 0.083 Fst = 0.04338, ). A. sachalinsis (D = 0.008) , A. nephrolepis (D = 0.005). . ., [33] 20 A. sibirica
: Fis Fit , 0.0432 0.0844, . 16
. 4.32%, 8.44%
Fst 0.431. D 0.0030. D , . 20 , 11 . . . . [32], . 7.33%-
(Fis =0.0733) 10,97%- (Fit =0,1097)- . 3,94% . , D 0,0041. . . [22] (A. nephrolepsis) 0.008, (A. sachalinensis) – 0.005. 0.047.
(GST = 0.082)
, Abies (GST = 0.063) [41].
2.2 AFLP – Abies
. AFLPs , . AFLPs - 17
[40]. AFLP , , , [43]. . . . . [25] AFLP- 20 Abies sibirica ( A. nephrolepis A. sachalinensis). 4 . , 168 117 . , A. sibirica, A. nephrolepis A. sachalinensis, (FCT = 0,53),
(FSC = 0,125). , A. sibirica AFLP – 14% (FST = 0,141). , , Abies sibirica. , . , Abies ziyuanensis (L.K.Fu & S.L.Mo), [54]. Abies, , A.ziyuanensis . , 139 , . A. ziyuanenesis He= 0,136, , . Gst = 0,482 (AFLP) . AMOVA (Φst = 0,550) (Φct = 0,139). , 18
, , , , , A. ziyuanensis.
2.3 Abies, SSR
. . Pinaceae, Abies, [47]. Abies 36 Abies, . . 5226 . , , . - , – . 39 , 23 , 16 – , . Abies ( ) 25. Abies 100 0.75 [26].
2.4 Abies, SSR
. . Pinaceae, Abies, [34].
19
. . . . [27] : A. sibirica, A. nephrolepis A. sachalinensis.
A. sibirica : No = 12.1, NE = 7.22,
HE = 0.8742.
A. nephrolepis : No = 17.3, NE =
11.9, HE = 0.9395.
A. sachalinensis : No = 15, NE=2.4,
HE=0.9637. A. sachalinensis 1,68% , A. sibirica (29 ) A. nephrolepis ( ) RST 8,34 14,83%, . Abies, 11,1% , 6,59% .
2.5 Abies, SSR
(Abies alba Mill) – , , . Abies alba. 5,2 , 0,532 24 Abies alba [37]. Abies alba 8 я я , я . я я . я 0,451 0,902 0,753 [38]. Abies nordmanniana (Stev.) Spach [42] .
20
,
(HE) 0,904.
2.6 Abies, ISSR-
ISSR- , [12], , - [8], , [57]. (Abies nephrolepis Max.) [58], - , - . 24 ISSR 248 , .
(HE = 0,240) , .
(GST = 0,082). (D = 0,027) . - . , - . , . . , , [2].
21
3
3.1 (SSRs)
5-50 . , . , , , . . , , . , . , , , . , . Abies sibirica Ledeb., [49]. CLC Assembly Cell. , , GMATo V1.2 [56]. 22
. 264864 , SSR- . , -, -, - ( 3). - WebSat [46], 140-280 . . , - . BLAST A. sibirica. , , ( 1). 128 .
1 — , A. sibirica
15 187 40 19 10 109 25 13 6 74 32 23 7 105 18 9 ∑ = 475 ∑ = 115 ∑ = 64
Abies sibirica « » .
23
3.2
. , , . A. sibirica, ( ) . , 8 - . «touchdown» . , . 100 - 200 , 30 ( 240 ). ( - ) [39]. : 94° — 1 ; 9 , 30 94° , 30 60° , 1 ° ( 50 ° ) 1 72° . 24 : 94° – 30 , 50 ° – 30 . 72 ° – 30 . 72 ° 10 . – ( 4 ° ).
24
«GenePak PCR Core» « », « » Taq- - , . 6%- -EDTA- . . pBR322 E. coli, HpaII.
3.3
8 , . 240, – 30. , 2.
2 – , № , \
58°21'36" . . 1 97°30'0" . . 419 58°19'48" . . 2 94°56'24" . . 293
25
56°57'0" . . 3 91°33'0" . . 232
55°40'48" . . 4 93°0'36" . . 333 55°18'36" . . 5 96°43'12" . . 423 2 54°23'24" . . 6 93°33'0" . . 542 52°57'36" . . 7 91°43'12" . . 642 52°48'36" . . 8 90°46'12" . . . 849
4 –
3.4
26
, (A), (Ho)
(He), (NE).
(NA) .
(Ho) .
= 1 2, (1)
fk — − , k- .
1 n = 1 , (2) − He — . F- [59].
(FIS)
= 1 (3) − Ho — ; — . (F ) IT
= + 1 , (4)
−
27
(FST) i- , ,
= , = 1 2 (5) − − — i- .
, , . . [48]
D = ln , = , (6) ( 2 )(푌 2 ) 푁 푁 − I — ;
jk jk — j- k- .
null- MICRO- CHECKER [55]. null- , . GenAlEx 6.5 [51]. - (UPGMA) [52] STATISTICA [53]. [45].
28
4
4.1
, 64 15 , 10 , 15 . 24 - . ( A). 10 , ( 3).
3 – , Abies sibirica
29
11 .
30
А 10 , - - , - . 26 . 0 4.
(NA = 3,2; NE = 2,52; HO = 0,550; HE =
0,516) (NA = 3,1; NE = 2,37; HO = 0,627;
HE = 0,540). Abies A. sibirica , 9,1%
(FIS =-0,091) 5%-
(FIT= -0,045) . 5%
(FST = 0,045) . . ,
. (DN) ( 0,014 0,108). , ( , , , , ) ,
FST = 0,037; DN = 0,077. , ,
31
: - .
32
А
1. , . . / . . , . . , . . // . 1983. – . 19. – № 2. – . 264–276. 2. , . . / . . , . . // . – 2002. – . 38, № 9. – . 1173–1195. 3. , . . : , , [ ] / . . , . . , . . , . . . – : http://www.agroatlas.ru.
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34
20. , . . (Abies sibirica Ledeb.) / . . // . . . . – 1971. – . 105–111. 21. , . . (Pinus sibirica Du Tour) / . . // . – 2014. – . 50, № 9. – . 1059–1059. 22. , . . : . . … . . : 03.00.05 / . – , 2004. – 38 . 23. , . . / . . . - , 1949. – №5. – 86 . 24. [ ] : . – : http://www.pro-rasteniya.ru/. 25. , . . (Abies sibirica Ledeb.), AFLP- / C. . , . . // . – 2011. – . 47, № 2. – . 272-278. 26. , . . Abies / . . , . . // . – 2014. – . 50, № 4. – . 420–420. 27. , . . (Abies sibirica Ledeb.) A. nephrolepis (Trautv.) Maxim. A. sachalinensis Fr. Schmidt / . . , . . // . – 2007. – . 43, № 12. – . 1637–1646. 28. , . . - (Abies Mill., Pinaceae) - : . . … . . : 03.00.05 / . – , 2008. – 24 .
35
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33. , . . - (Abies sibirica Ledeb.) : . . … . . : 03.00.05 / . – , 2006. – 17 .
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39
А
, Abies sibirica
.1 - , c F:CTCAAAGGTAGTGTTTCTCAGTGGTAT As_43741 (TCA)15 219 R:CAATGATGACAATGATGTTGGC F:GGGAGATTGATTATGTGGATCA As_99437 (ATG)17 228 R:CATGCAAGGGGACTAATATCTTC F:ACGAAGGTGAAGGGTTCTATCA As_279869 (TTA)15 253 R:TTAAGGGGAATTGTTGTGGGTA F:CAATGATGATGGACTAGATGGA As_301948 (ATC)16 214 R:AAGTCTGATGGGTTATGCTTGT F:TTACCCTCGAATCTAAAGGTGG As_878430 (AAT)15 260 R:CTATCCCACAAACAAAGGGGT F:GTGATCTATGCCTCTCCAATGA As_1177607 (TAA)17 188 R:ACTGGTCACAATGGTTTGAATG F:GTGAGGAGGAGATTTGTAAGTGC As_1563074 (ATT)18 142 R:CAGATGAGTACGAACCACTACCC F:CCTTCCACCTTCCATGATATTC As_420232 (TAAA)10 182 R:TCCAAGAAAAGGCACAAGAAGT F:AAGTTGTTTGATCCCTCTCGAC As_563614 (ATAG)12 157 R:CATGTAATGCTCCAATCAGGAA F:CCCTTACCTCCCTAACACTTGA As_721750 (TATG)10 147 R:GGCAAGATAGATGATTGACACC F:CACTAAAAGAGGAGCCATGTCC As_2280886 (TGTA)10 201 R:GACACCTGCAATCACCAAGAC F:GATGAGAGATGCTTGTGAATCC As_2687764 (TATG)10 222 R:GGAGGTTATCATGGAAGTGGAC As_2996373 (ATAC)11 F:TCGCTTCCACACTGACCAC 272
40
R:TTACACACAAAAGGGCTCACAC F:GTCGCAGAATGTTGAATTGCT As_3602348 (CATA)10 278 R:CATGGATGGAGAGGACAAGTTT F:AAGGTTCTTAAATTCCAACCCC As_347605 (GAAAA)6 253 R:GACTTGAGGATGAGGTTCTTGC F:GGATGCAGGGTTTCATTTATTC As_1133359 (GATTT)6 238 R:CCAGAAGATTCGACATGGAAA F:TAGACTTTGAGGTGATGGGCTT As_1356647 (GATAA)7 250 R:TTGCTCCGTGATCTTAGGTTTT
.1 F:TGTTTTCCAGATTTGTGAGTGC As_2288295 (AGATA)6 172 R:AGTCCCGTTGACCTACTTTGTG F:CTCCCACATCAATCCCATTTAC As_3458593 (TTATC)7 225 R:TGGTTTCCTGACGATGACTCTA F:TAACGACCTGCTGCTATTTTCA As_404145 (CCAGCT)8 187 R:GGGCACCATTCTCAATTTCTAC F:ATGCTGCCCATTGAGAATTAGT As_1898225 (TCTTCA)9 270 R:ACCGCTCTTCCCTTTACTTTCT F:TTTGGACTGAGATTTGGGC As_1970306 (GCAAGA)7 248 R:CGATTATGAAAGAATGGGTAGC F:CCCATCCTATCTCCTACACTTCA As_2631734 (CACTCC)7 169 R:GGGTTGATATGTGAAAAGGGAA F:CGTTCTTTCACTTTTGTTGGC As_3666591 (TATTCC)7 258 R:TCCTTTGCCATTGTTCATGT
41