,2017,37(10): 犃犮狋犪犅狅狋. 犅狅狉犲犪犾. 犗犮犮犻犱犲狀狋. 犛犻狀. : (2017) 犱狅犻 : /j.issn "#$%&' ( )* "#+,-./0 1,2,

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1 ,2017,37(10): 犃犮狋犪犅狅狋. 犅狅狉犲犪犾. 犗犮犮犻犱犲狀狋. 犛犻狀. : (2017) 犱狅犻 : /j.issn "#$%&' ( )* "#+,-./0 1,2, 1, 1, 1, 1,2, 1,2 ( :,; ; #$,; ) 1 : <=>(GISH)?@ABCD E ) = F G ) H ;,I J K L * MNO O,#$ IP F Q R S < = > T U E F < = > V W <=>HX <=>S Y F Z [ \ ] ^ _ ` Z *ab c^v H=> ^ N G < = \ M 8 9 #$ Q, E 8 9 D, G / E F, \ # $ 8 9 ) B \ +, C D S?@ : <=>;89;E = CD; ); F QR - 2:Q :A 犚犲狊犲犪狉犮犺犘狉狅犵狉犲狊狊狅犳犌犲狀狅犿犻犮犻狀狊犻狋狌犎狔犫狉犻犱犻狕犪狋犻狅狀犪狀犱狋犺犲犃狆狆犾犻犮犪狋犻狅狀犻狀犎狅狉狋犻犮狌犾狋狌狉犪犾犘犾犪狀狋狊 WANG Yan 1,2,CHEN Qing 1,CHEN Tao 1,ZHANGJing 1, TANG Haoru 1,2,WANG Xiaorong 1,2 (1ColegeofHorticulture,SichuanAgriculturalUniversity,Chengdu611130,China;2InstituteofPomologyandOlericulture, SichuanAgriculturalUniversity,Chengdu611130,China) 犃犫狊狋狉犪犮狋 :Genomic 犻狀狊犻狋狌 hybridization (GISH)hasbeenusedtodiscriminatethediferentancestralge nomedonorsinpolyploidsorhybrids,analyzethephylogeneticrelationshipsbetweencultivarandwildrel atives,andstudychromosomebehaviorofmeiosisinplants.gishtechnologyincludesmulti colorgenomic 犻狀狊犻狋狌 hybridization(mc GISH),comparativegenomic 犻狀狊犻狋狌 hybridization(cgish),andself genomic 犻狀狊犻狋狌 hybridization(self GISH).Itskeyprocedurescontainchromosomepreparation,probepreparationand lengthoptimization,theconcentrationratioofprobetoblockingdna,andstringencyofpost hybridiza tionwashes.thispaperreviewedtheresearchprogressonthedevelopmentofgishanditsutilizationin horticulturalplants,andsummarizedtheirpromisingapplicationprospects.duringthepastseveralyears, thecompletegenomesequencesareavailableforhorticulturalplantssuchasgrape,banana,pear,sweet orange,andcucumber.itisnecessaryforresearcherstoselectmorechromosomalspecificmarkersfrom thewholegenomicsequence.combing withfluorescencebandingandfluorescence 犻狀狊犻狋狌 hybridization : ; : : ( , )!: (1987-),,,!"#$%,&' ()*+,- #$ E mail:wangyanwxy@163.com./01:,, 2,&' ()*+,- #$ E mail:wangxr@sicau.edu.cn

2 (FISH)technologies,GISH wilbecomeapowerfultooltofurtherilustratetheoriginandidentifygenetic relationshipofhorticulturalplants. 犓犲狔狑狅狉犱狊 :genomic 犻狀狊犻狋狌 hybridization (GISH);horticulturalplants;polyploidorhybrididentification; speciesorigin;chromosomebehavior < = > (genomic 犻狀狊犻狋狌 hybridiza tion,gish) < = > (fluorescence 犻狀狊犻狋狌 hybridization,fish)?@ G [1],&' " / DNA 0R Z, / DNA 0Rab, * F Q <=>; 12 I, ab, =>, I G F GISH?@ E = C D O * `S#$, I +, ' < [2 3] GISH?@ Y I B \ 8 9 #$,B R89 I - +, 1 <=>?@ Y <=>?@IR5 &' : Z [H F [ => F\ IJ M, F Z = >H=> S R Y :; GISH = >, F ' H, F, Z DNA A\ Q F A, WR,! "B#$ [4] %& ' ( 犚狌犫狌狊 ),A [5] [ F,=> 2% ( ) *! H +!, - 37."1.5h,A [6] =>/ 0, F '1,B 2 H / 0 F 6 7 Hes lop Harrison [7], F 2 H=>, 4% (w/v)e 8 9 : (paraformaldehyde) D 10 min, ;, < F =, A >=>? 1.2 : < DNA ABC,DE ; Z?A, DNA B ^' E F G H I,OD 260 /OD 280 R 1.8~1.9 DNA JK L,GISH DNA Z]^M D ', N]^R300 ~500bp [8] ZO => D,O] (>1kb)D PQ F R,ST=> [9] 1.3 =>? 犇犖犃.@ AB GISH UV,a b DNA _ W * X Y = >, DNA ZD* Y =>, Z*ab DNA c^v ' Z*a b V U O Z N [ W7, O AN, ' ab?a7; O AZ, ' ab?a,\]a B ab [10] ^ _ ( 犘狅狀犮犻狉狌狊 )* ` a ( 犆犻狋 狉狌狊 ) O WZ, Z*ab DNA c^v R1 4 b Ic F [11] ' = CD, Z*abV R1 50,J IcM ( 犚. 狆犪狉狏犻犳狅犾犻狌狊 )H ( 犚. 犮狅狉犲犪狀狌狊 ) [6] 1.4 &'CDE => ^H ^M D GISH / Y ^AB =>I D =>I, &' ^ c^\ G D ^ =>,= > DNA F I ^ AB : ^ (%)=100 (%) - 犉 m( 犜 m- 犜 h) [12], 犜 m = I ^; 犜 h = I. ^; 犉 m,* Z]^ ( Z>150bp, 犉 m = 1; Z<50bp, 犉 m=5) #$,=> ^A./ H9 c^b\=> ^G 2, 20%9 2 SSC min,GISH => ^A 80%B [10] 2 GISH?@ I "#$%&' *(biotin)h (digoxygenin) G) DNA, )E HNO =

3 10,S: 89 #$ 2089 QEF < = > (multi colorgenomic 犻狀狊犻狋狌犺狔犫狉犻犱犻狕犪狋犻狅狀, 犿犮 犌犐犛犎 ),B E C D [2] mc GISH D ) => < F G) V X GISH I ' ; ( 犕狌狊犪 ) [13 15] ( 犇犻狅狊狆狔狉狅狊 ) [16] H ( 犛狅犾犪 狀狌犿 ) [17] S89 G)H ;#$ 2.2 A "#$%&' B DNA R Z,* M F Q=>, *M 4Y ), ` 4 Y F I, O,b R V W < = > (com parativegenomic 犻狀狊犻狋狌 hybridization,cgish) cgish?@ E 8 9 \ M N O O #$,R )\ ` I [18 25] 2.3 F "#$%&' B DNA R Z,* X F Q= >,b R X < = > (self ge nomic 犻狀狊犻狋狌 hybridization,self GISH ) self GISH / 3 4 DNA Y F =>, 4Y H F I H #$,self GISH / [, * 7 [6,26 27], self GISH [ AB 2 [28 30] 3 GISH 89 #$,GISH?@ (Musaceae) %& (Rosaceae) (Rutaceae) (Crucife rae) (Solanaceae)H (Liliaceae)S89 #$,&' E ) = F G) KL * W O IP F QRS#$ 3.1 G9*HIJ, #$,./ IP/0 F QRG I )* )E 7 E, L ;*G),./ GISH DABL E ) ( 1) ^ * S [31] ( 犆犻狋 狉狌狊犵狉犪狀犱犻狊 cv. Shatianyou ) 3 H 3 Q GISH IJ, 3 ) M )3,E * 3 R )3 ; )3./ =>, B NO 犐狆狅犿狅犲犪狋狉犻犳犻犱犪 (2 狓 ) R Z,* 犐. 狋狉犻犳犻犱犪 (4 狓 )2 F Q GISH #$,, F! " /,A # 犐. 狋狉犻犳犻犱犪 $% G; & ' F M =>/, / *"^ [,A F H I2 [32] ( 犆犪犿犲犾犻犪狉犲狋犻犮狌犾犪 狋犪 )E 4 ) #$,LiuS [33] B NO ( 犆. 狆犻狋犪狉犱犻 )H ( ) * ( 犆. 狊犪犾狌犲狀犲狀狊犻狊 )0R Z,B# 0Rab,GISH, *3 H+ =>/, ()* *+ =>/,, )3 # H =>-, )+ D )3 *()* => G ).\ 2 \B )E, F EF GISH 77 M ; / Lim S [34] B 犐狉犻狊狊犲狋狅狊犪 (2 狀 =38) H 犐. 狏犻狉犵犻狀犻犮犪 (2 狀 =70) R Z,* 2 F 0 1 ( 犐. 狏犲狉狊犻犮狅犾狅狉,2 狀 =108) F = >, I 38 H70 F, 2F0 1R ) E I B 犛狅犾犪狀狌犿狏犲狉狉狌犮狅狊狌犿 (AA) 犛. 犼犪犿犲狊犻 (BB)H 犛. 犪狀犱狉犲犪狀狌犿 (PP)R Z,* ( 犛. 犺狅狌犵犪狊犻,2 狀 =6 狓 =72) H 犛. 犱犲犿犻狊犻狌犿 (2 狀 =6 狓 =72) F =>, 犛. 犺狅狌犵犪狊犻 R )+,A B H P * M ;; 犛. 犱犲犿犻狊狊犻狌犿 D R A ; )+ [17] 3.2 &HKLMN OJ E KL,- R I,. => S 23= 4>, B 5 +, 6 KL <=>?@ 7 F => 8,9\ : [2], 8 9= Q GISH IJ, I F G),R- 0 I +, GISH 89= F G)H ;C D ; 1 LandiS [35] FISH H GISH

4 犌犐犛犎 ()* G9 &H789# MN-./0 Table1 IdentificationofchromosomecompositioninpolyploidsandhybridsrevealedbyGISHinhorticulturalplants Plantspecies F number F ; composition GISH GISHresult Reference P Q 犚狅狊犪犮犲犪犲 Naturaltriploid 2 狀 = 3 狓 = 51 (3 # ) 犈狉犻狅犫狅狋狉狔犪犼犪狆狅狀犻犮犪 (tetra 2 狀 = 3 狓 = 51 ploid diploid) T U ) H ) Naturaltriploids containedbothautotriploidsandalotriploids. 34 F G 3,17 G # 34chro mosomeswerefromtetraploidand17fromdiploid. <= >?= 犉狉犪犵犪狉犻犪狀犻犾犵犲狉狉犲狀狊犻狊 犉. 犪狀犪狀犪狊犪 2 狀 = 5 狓 =35 F n F a F a F a F a 7F n +28F a [39] 犉. 犪狀犪狀犪狊狊犪 ( 犉. 狏犲狊犮犪 犉. 犻狀狌犿犪犲 ) 2 狀 = 8 狓 = 56 F v F v F i F i F i F i F i F i 14F v +42F i [40] [37] A F B C 犘狉狌狀狌狊犮犲狉犪狊狌狊 Schatenmorele 2 狀 = 4 狓 = 32 ' 犚狌犫狌狊 L. 16 F A D B C,16 G= BC 16 chromosomeswerefrom 犘. 犪狏犻狌犿 and16from 犘. 犳狉狌狋犻犮狅狊犪. [41] 2 狀 = 4 狓 = 28 R R BB 14R +14B [36] 2 狀 = 5 狓 = 35 R R R BB 22R +13B Aurora 2 狀 = 8 狓 = 58 R BBBBBBB 9R +49B E = Putativehybrid 2 狀 = 3 狓 = 21 R S Q 犚狌狋犪犮犲犪犲 14 F G =,7 G 14chromosomes werefrom 犚. 狆犪狉狏犻犳狅犾犻狌狊 and7from 犚. 犮狅狉犲犪狀狌狊. _ 犘狅狀犮犻狉狌狊狋狉犻犳狅犾犻犪狋犪 犆犻狋狉狌狊犪狌狉犪狀狋犻狌犿 2 狀 =4 狓 = 36 P t P t C a C a 18P t +18C a [42] _ 犘. 狋狉犻犳狅犾犻犪狋犪 犆. 狊犻狀犲狀狊犻狊 2 狀 =2 狓 =18 P t C s 9P t +9C s [11] _ 犘. 狋狉犻犳狅犾犻犪狋犪 犆. 狆犪狉犪犱犻狊犻狊 2 狀 =2 狓 =18 P t C p 9P t +9C p 犆. 犵狉犪狀犱犻狊 cv.shatianyou 2 狀 = 4 狓 = 36 犆. 犵狉犪狀犱犻狊 cv.shatianyou 犆. 犻犮犺犪狀犵犲狀狊犻狊 3 ER )3, 3 R ) 3 Naturaltetraploids were mainly alotetraploid, whileartificialtetraploidswerealautotetraploid. [6] [31] 2 狀 = 2 狓 = 18 C g C i 9C g +9C i [43] 2 狀 = 4 狓 = 36 C g C g C i C i 18C g +18C i 犆犻狋狉狅狆狊犻狊狊犮犺狑犲犻狀犳狌狉狋犺犻 Nanputangor( 犆犻狋狉狌狊 spp.) 2 狀 = 2 狓 = 18 C s C n 9C s +9C n [44] 犆. 狊犻狀犲狀狊犻狊 cv.hongjiangcheng 2 狀 = 4 狓 = 36 ) 3 Naturalautotetraploid [45] T U V 犕狌狊犪犔. Calcuta4,TMP2 狓 狀 = 2 狓 = 22 AA 22A Butohan2 2 狀 = 2 狓 = 22 BB 22B Kunnan,SafetVechi 2 狀 = 2 狓 = 22 AB 11A+11B Wompa 2 狀 = 2 狓 = 22 AS 11A+11S Bluggoe,Fougamou,Praha 2 狀 = 3 狓 = 33 ABB 11A+22B Pelipita 2 狀 = 3 狓 = 33 ABB 8A+25B Kunnan4 狓 Cameroun 2 狀 = 3 狓 = 33 ABB 11/10A+22/23B Agbagba,FiguePomme,Laknao,Malaccencis Kunnan 2 狀 = 3 狓 = 33 AAB 22A+11B 4 狓,Mbi,ObinoI Ewai,P.rajabulu Luba 2 狀 = 3 狓 = 33 AAB 22/23A+11/10B M.bouroukou,Nyombe 2 狀 = 3 狓 = 33 AAB 21/22A+12/11B Kunnan4 狓 IDN110 2 狀 = 3 狓 = 33 AAB 23/22/21/20A+10/11/12/13B Karoina 2 狀 = 3 狓 = 33 AAT/ATT 22A+10T Kunnan4 狓 2 狀 = 4 狓 = 44 AABB 22A+22B TMPx 狀 = 4 狓 = 44 AAAB 33A+11B Yawa2 2 狀 = 4 狓 = 44 ABBT 11A+22B+10T W X Y V 犘犪狊狊犻犳犾狅狉犪 [13 15, 46 47] HD ( 犘. 犵犪狉犱犻狀犲狉犻 犘. 犵犻犫犲狉狋犻 ) 2 狀 = 2 狓 = 18 HD ( 犘. 狊狌犫犾犪狀犮犲狅犾犪狋犪 犘. 犳狅犲狋犻犱犪 ) 2 狀 = 2 狓 = 22 P s P f 10P s +12P f HD ( 犘. 狊狌犫犾犪狀犮犲狅犾犪狋犪 HD13 133) 2 狀 = 2 狓 = 22 P s H 8P s +14H HD ( 犘. 狊狌犫犾犪狀犮犲狅犾犪狋犪 HD13 133) 2 狀 = 2 狓 = 22 P s H 9P s +13H 9 F G 犘. 犵犪狉犱犻狀犲狉犻,9 G 犘. 犵犻犫犲狉狋犻 9chromosomeswerefrom 犘. 犵犪狉犱犻狀犲狉犻 and9from 犘. 犵犻犫犲狉狋犻. 犘. 狊狌犫犾犪狀犮犲狅犾犪狋犪 犘. 犳狅犲狋犻犱犪 2 狀 = 2 狓 =20-22 P s P f 9-16P s +6-11P f [49] [48]

5 10,S: 89 #$ ContinuedTable1 Plantspecies F number F ; composition GISH GISHresults References 2 狀 = 3 狓 = 30 P s P f 21P s +9P f V 犇犻狅狊狆狔狉狅狊 犇. 犽犪犽犻 犇. 犵犾犪狀犱狌犾狅狊犪 2 狀 = 8 狓 = 120 D k D k D k D k D k D k D g D g 90D k +30D g [16] V 犣犻狕犻狆犺狌狊 F G 7 H 犣. 犼狌犼狌犫犪 Zanhuangdazao 2 狀 =3 狓 =36 ) Autotriploid [50] Q 犆狉狌犮犻犳犲狉犪犲 犅狉犪狊犻犮犪狀犪狆狌狊 (AACC) 犚犪狆犺犪狀狌狊狉犪狆犺犪狀犻狊狋狉狌犿 (RR) 2 狀 =2 狓 =28 19AC+9R [51] 犅. 狀犪狆狌狊 (AACC) 犅. 狅犾犲狉犪犮犲犪 (CC) 2 狀 =3 狓 =27 ACC 9A+18C [52] 犅. 狉犪狆犪 ssp. 狆犲犽犻狀犲狀狊犻狊 (AA) 犅. 狅犾犲狉犪犮犲犪 var. 犮犪狆犻狋犪狋犪 (CC) 2n=17-20,26 Q 犛狅犾犪狀犪犮犲犪犲 AC 7A+10C,9A +9C,10A +9C,12A +8C,14A + 12C 犛狅犾犪狀狌犿犺狅狌犵犪狊犻 2 狀 =6 狓 =72 AABBPP ) + Alohexaploid:24A+24B+24P [17] 犛. 犱犲犿犻狊狊狌犿 2 狀 =6 狓 =72 AAAAAA ) + Autohexaploid:72A [53] I 犔. 犲狊犮狌犾犲狀狋狌犿 犛. 犾狔犮狅狆犲狉狊犻犮狅犻犱犲狊 2 狀 =3 狓 +2=38 L e L e S s F G 犔. 犲狊犮狌犾犲狀狋狌犿,12 G 犛. 狊犻狋犻犲狀狊,2 G 犛. 犾狔犮狅狆犲狉狊犻犮狅犻犱犲狊 24chromosomes werefrom 犔. 犲狊犮狌犾犲狀狋狌犿 and12from 犛. 狊犻狋犻犲狀狊 and 2from 犛. 犾狔犮狅狆犲狉狊犻犮狅犻犱犲狊. [54] I 犔. 犲狊犮狌犾犲狀狋狌犿 犛. 狊犻狋犻犲狀狊 2 狀 =2 狓 +1=25 L e S s +1 Z V 犃犾犻狌犿 12 F G 犔. 犲狊犮狌犾犲狀狋狌犿,12 G 犛. 狊犻狋犻犲狀狊,1 G 犛. 犾狔犮狅狆犲狉狊犻犮狅犻犱犲狊 12chromosomes werefrom 犔. 犲狊犮狌犾犲狀狋狌犿 and 12 from 犛. 狊犻狋犻犲狀狊 and1from 犛. 犾狔犮狅狆犲狉狊犻犮狅犻犱犲狊. 犃. 犮犲狆犪 犃. 狉狅狔犾犲犻 2 狀 =40 犃. 犳犻狊狌狋狌犾狅狊狌犿 犃. 狋狌犫犲狉狅狊狌犿 2 狀 =3 狓 =24 20 F G 犃. 犮犲狆犪,20 G 犃. 狉狅狔犾犲犻 20chromo someswerefrom 犃. 犮犲狆犪 and20from 犃. 狉狅狔犾犲犻. 8 F G 犃. 犳犻狊狌狋狌犾狅狊狌犿,16 G 犃. 狋狌 犫犲狉狅狊狌犿.8chromosomeswerefrom 犃. 犳犻狊狌狋狌犾狅狊狌犿 and16from 犃. 狋狌犫犲狉狅狊狌犿. [55] [56] 犃. 狑犪犽犲犵犻 2 狀 =2 狓 =16 [ Q 犆狅犿狆狅狊犻狋犪犲 8 F G 犃. 犳犻狊狌狋狌犾狅狊狌犿,8 G 犃. 犮犲狆犪. 8chromosomeswerefrom 犃. 犳犻狊狌狋狌犾狅狊狌犿 and8from 犃. 犮犲狆犪. 7 J K L M L 犇犲狀犱狉犪狀狋犺犲犿犪犮狉犪狊狊狌犿 犆狉狅狊狊狅狊狋犲狆犺犻狌犿犮犺犻狀犲狀狊犲 2 狀 =6 狓 =54 D c D c D c D c D c C c 45D c +9C c [57] N L M L 犇. 犻狀犱犻犮狌犿 犆. 犮犺犻狀犲狀狊犲 2 狀 =3 狓 =27 D i D i C c 18D i +9C c [58] LineT3 2 狀 =6 狓 =54 \ ] ^ V 犅犲犵狅狀犻犪 =,9 F G M L,9 G O,36 G 犆. 犵狉犪狀犱犻犳犾狅狉狌犿 H 犆. 犮狉犪狊狊狌犿.Atruetri generichybrid containing 9 chromosomesfrom 犆. 犮犺犻狀犲狀狊犲,9from 犃狉狋犲犿犻狊犻犪狏狌犾犵犪狉犻狊,and36from 犆. 犵狉犪狀犱犻犳犾狅狉狌犿 and 犆. 犮狉犪狊狊狌犿. 犅. 狊狅犮狅狋狉犪狀犪 2 狀 =2 狓 =28 SS 28S [60] Tuberousbegonia 2 狀 =4 狓 =56 TTTT 56T Elatior begoniashybrids 2 狀 =2 狓 =28 TT SS 14T+14S _ ` V 犔犻犾犻狌犿 2 狀 =3 狓 =42 TTTT SS 28T+14S 2 狀 =3 狓 =41 TTTT SS 27T+14S 2 狀 =4 狓 =56 TTS SSS 28T+28S MarcoPolo 犔. 犺犲狀狉狔犻,Expression 犔. 犺犲狀狉狔犻 2 狀 =2 狓 =24 12 F G MarcoPolo (Expression),12 G 12chromosomeswerefrom MarcoPolo (Epression)and12from 犔. 犺犲狀狉狔犻. 犔. 犾狅狀犵犻犳犾狅狉狌犿 Asiatichybrid (LAhybrid) 2 狀 =2 狓 =24 LA 12L+12A [62] a S V 犜狌犾犻狆犪 F1 ( 犜. 犵犲狊狀犲狉犻犪狀犪 犜. 犳狅狊狋犲狉犻犪狀犪 ) 2 狀 =2 狓 =24 T g T g T f T f 12T g +12T f [63] BC1 ( 犜. 犵犲狊狀犲狉犻犪狀犪 F1 ) 2 狀 =2 狓 = T g +6-4T f 2 狀 =4 狓 =48 42T g +6T f BC2 ( 犜. 犵犲狊狀犲狉犻犪狀犪 BC1 ) 2 狀 =2 狓 = T g +3-0T f 2 狀 =2 狓 +1=25 25T g [59] [61]

6 ContinuedTable1 Plantspecies F number F ; composition GISH GISHresults References b c V 犃犾狅犲 犃. 犪狉犫狅狉犲狊犮犲狀狊 犃. 狏犲狉犪 2n=2x=14 犃狊犮狅犮犲狀犱犪 犘犺犪犾犪犲狀狅狆狊犻狊 2 狀 =3 狓 =62-67 #,7 F G7,7 G Alo diploid:7chromosomeswerederivedfrom 犃. 犪狉犫狅狉犲狊 犮犲狀狊 andtheremaining7originatedfrom 犃. 狏犲狉犪. V 犖犪狉犮犻狊狊狌狊 [64] 28A+34P,31A+36P [65 ] F 犖. 狋犪狕犲狋犪 var. 犮犺犻狀犲狀狊犻狊 two colorflower 2 狀 =3 狓 =32 V 犐狉犻狊犔. H H = Naturaltriploid hybrid between white flower narcissus (2 狀 )and yelowflowernarcissus( 狀 ). 2 F 0 1 犐. 狏犲狉狊犻犮狅犾狅狉 2 狀 =108 := > R ) E,38 F G 0 1,70 F G 犐. 狏犻狉犵犻狀犻犮犪 Alopolyploid:38 chromosomes werefrom 犐. 狊犲狋狅狊犪 and70from 犐. 狏犻狉犵犻狀犻犮犪. [66] [34] Note:Thehybridizationcrossesareal maternalparent paternalparent 1 # P H1 3 Q P R S Q # $, GISH - ; FISH T Z EUHE <V 7? F,, * B \ E ] L W R S D Lim S [36] KL X(2 狓 ) Q K L X(2 狓 ~12 狓 )B\ ) E Q GISH I J, )Y 3 F G (raspberry),2 G Q (blackberry); Q ) E ^ Q K L P Au rora F F,M < F, R ' # =>H ( 犕狌狊犪 L.)K L I A B SH T S4 F ;, I R 犕. 犪犮狌犿犻狀犪狋犪 犕. 犫犪犾犫犻狊犻犪狀犪 犕. 狊犮犺犻狕狅犮犪狉狆犪 H 犃狌狊狋狉犪犾犻犿狌狊犪 E < 1 GISH L CD E KL G )H ; [13 15,46 47], = Pelipita (2 狀 = 3 狓 = 33,ABB) F ;R8A+25B Z 11A+22B KarlovS [67] 3 O([ ) A(\ A ) L(] ) I c G NathewetS [68] GISH, ^_= ( 犉. 犻犿狌犿犪犲 )H`a= ( 犉. 狏犲狊犮 a)a * K L = ( 犉. 犪狀犪狀犪狊狊犪 Hokowase,2 狀 =8 狓 =56) ; GISH M 89ZO= L-CD b '0,CD = c, O B R,Singh S [51] 5 ( 犅狉犪狊犻犮犪狀犪狆狌狊,2 狀 =38,AACC)* ( 犚犪狆犺犪狀狌狊狉犪狆犺犪狀犻狊狋狉狌犿,2 狀 =18,RR)= >, GISH M I P F Q R D, AB 0 6 GISH M 7H S = >H [52,69] TangS [57]./ GISH 7J KL( 犇犲狀犱狉犪狀狋犺犲犿犪犮狉犪狊狌犿,2 狀 =10 狓 =90)H M L ( 犆狉狅狊狊狅狊狋犲狆犺犻狌犿犮犺犻狀犲狀狊犲,2 狀 =2 狓 =18) = > 8 (2 狀 =6 狓 =54)c Q C D, M O,GISH?@ E;R89 2 H P L-CD ': 3.3 *H K O # $ = > - " 89KL. E W =>E,M+, 4=, GISH,AL ' O, A # $' )* `, R89 () " ChoiS [19] B # ( 犇. 犵犾犪狀犱狌犾狅狊犪犻 ) DNA R Z, KL ( 犇. 犽犪犽犻,2 狀 = 6 狓 = 90) F H. W /, R # *KL WN O,, MA KL W GISH K L ( 犕. 犻狀犱犻犮犪 )* a ( 犕. 狊狔犾狏犪狋犻犮犪 ): W N O [20] Preeda sutijits [70] GISH IJ ( 犆. 狌狀 狊犺犻狌 )*_( 犘. 狋狉犻犳狅犾犻犪狋犪 ) WN O ;Kita jimas [71] *`a( 犆. 狉犲狋犻犮狌犾犪狋犪 ) MWN, ( 犆. 犿犪狓犻犿犪 )A * M ) NN S [50] # $ H ( 犣. 犼狌犼狌犫犪 )H F H ( 犣. 犼狌犼狌犫犪 subsp. 狊狆犻狀狅狊犪 ) : ^ ),R H ) FH 1 I +, Lee S [72]. / ( 犘犪狆犺犻狅狆犲犱犻犾狌犿 ) => 8 IP F

7 10,S: 89 #$ 2093 Q R GISH I J, O B R 8 9 ) `HI #$ ' , F QR* DNA 4 ` \] ; = GISH #$, ',G = 8 Q,' D, [1], = F,, [73] ^ K L Figuepomme (AAB)H Praha (ABB),B &' F., A F < D 2`, A F > I [47],M A * B A [74] ( 犛犪犮犮犺犪狉狌犿 spp.)* ( 犈狉犻犪狀狋犺狌狊犳狌犾狏狌狊 ) = > 8, +, +, 0 F \ ` F 8 P,, F B I I [75] DengS [76] KL L ( 犇. 犿狅狉犻犳狅犾犻狌犿 )* O ( 犃. 狏狌犾犵犪狉犻狊 ) =,G) F = : RI V, L H O WZ./ <=>, ) F G), CD F H < [2] ( 犔犻犾犻狌犿 ) ; F 1 8 E A * O,GISH 4 > 8 +, E U - S T [77] Nat enapits [78] LLO = H# \A Q=>, ) IP/0 IP ;H ) (ACC) I,^ Ⅰ F, Ⅰ Ⅱ / 0 F H = F, A * IP [52] JeridiS [46] D. / # $ 3 KL (AABB), ( 犕. 犪犮狌犿犻 狀犪狋犪,AA)H ( 犕. 犫犪犾犪犻狊犻犪狀犪,BB) > R )+, 4 W H S89, 7ERE, `0^, c, F E [79],./, IJC ZO= ) F M ", DNA R Z,*E = F Q <=>, CDE F ) IJ= F G), WR ' 2, W R ;, M F 2 W 2 Q GISH #$, $ 3 P O, A,GISH M M,^ I ' F # $ 5 (CMA + /DA PI - )H 4Y Z(^rDNA B\ Y S)\M I?@ G, E F ^ CMA + /DAPI - 5 F IR A~G S8, *rdna S 4 Y D <,A F [80] E 89 D, G / E F, <=>IJ,GISH M5 89 #$ R H 3: [1] SCHWARZACHER T,LEITCH A R,BENNETT M D, 犲狋犪犾. 犐狀狊犻狋狌 localizationofparentalgenomesina widehybrid [J]. 犃狀狀犪犾狊狅犳犅狅狋犪狀狔,1989,64(3): [2] CHESTER M,LEITCH A R,SOLTISPS, 犲狋犪犾.Reviewof theapplicationofmoderncytogeneticmethods(fish/gish) tothestudyofreticulation (polyploidy/hybridisation)[j]. 犌犲狀犲狊,2010,1(2): [3] RAINASN,RANIV.GISHtechnologyinplantgenomere search[m]//ksa,as(eds.)painting.neth erlands:kluweracademicpublishers,2001: [4],,. IP F 0 [J].,1979,21(3): CHENRY,SONG W Q,LIXL.Anewmethodofpreparing chromosomefrom plant[j]. 犃犮狋犪犅狅狋犪狀犻犮犪犛犻狀犻犮犪,1979,21 (3): [5],,,S. [ F #$[J].34567,2007,25(3): WANG X R,TANG H R,LIL, 犲狋犪犾.Studiesonroot tip squashing methodforchromosomepreparationof 犚狌犫狌狊 L. [J]. 犑狅狌狉狀犪犾狅犳犛犻犮犺狌犪狀犃犵狉犻犮狌犾狋狌狉犪犾犝狀犻狏犲狉狊犻狋狔,2007,25 (3): [6] WANG Y,WANG X R,CHEN Q, 犲狋犪犾.Phylogeneticin sightintosubgenera 犐犱犪犲狅犫犪狋狌狊 and 犕犪犾犪犮犺狅犫犪狋狌狊 ( 犚狌犫狌狊,

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