,2017,37(4): 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 "#$%& ' 1, 1, 1, 2 (1 345

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1 ,2017,37(4): 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 "#$%& ' 1, 1, 1, 2 ( , ; :8, ) ( ):; ; A B C D E (PPIase),! ' F G,H! I J K L MNIJO,! PQ ; RSNT U <,V=>SW; XY,!S) ' *Z[.\,]^_`abc X ' * + > ; UXY L%&, ; %& ^ *+,:; ; U;'Y -./0:Q789 1 :A 犚犲狊犲犪狉犮犺犘狉狅犵狉犲狊狅犳犘犾犪狀狋犆狔犮犾狅狆犺犻犾犻狀犌犲狀犲犉狌狀犮狋犻狅狀狊 TONG Huishan 1,WANGShanshan 1,ZHU Xinni 1,DING Wona 2 (1SchoolofMarineScience,NingboUniversity,Ningbo,Zhejiang315211,China;2ColegeofScienceandTechnology,Ningbo University,Ningbo,Zhejiang315211,China) 犃犫狊狋狉犪犮狋 :Cyclophilins (CyPs),membersofthe immunophilin proteinfamily,havepeptidyl prolylcis transisomeraseactivityandwidelydistributeindiferentorganisms.theyarehighlyconservedinprotein structureandexpressedincytosolandvariousorganeles.plantcypsaresupper familymembersandplay significantrolesinstressresponse,metabolicregulationandplantdevelopmentbesidestheircommonfunc tions.thisreviewsummarizesthefunctionsandrecentresearchprogressofplantcypsandaimstohelp theirfurthercharacterizationinthefuture. 犓犲狔狑狅狉犱狊 :cyclophilin;gene;biologicalfunction ; (cyclophilin,cyp) _ A(cyclosporineA,CsA) A B C D E (peptidyl prolyl 犮犻狊 狋狉犪狀狊 A X C KX L.\ [ X.\ ; FH! ', I L,,!IJK K IJ IJO [1] ; LT, ; =>SN PQ X PPIase,T ; V PPIase, > WD CCD RP RRM ZnFingerL TPR, ^ _ K K K.\ [2] ; RSN 5X <, > T X Y L 1990 Gas ser [3] ( 犔狔犮狅狆犲狉狊犻犮狅狀犲狊犮狌犾犲狀狋狌犿 ) ( 犣犲犪犿犪狔狊 ) ( 犅狉犪狊犻犮犪狀犪狆狌狊 ) L ;,!5, : ; : : (LY17C020002)!: (1992-),,!"#$%&', ()*+%& E mail:tonghuishan1@163.com,-./:01, $,, 2' E mail:dwn@zju.edu.cn

2 4, :; UY %& 831! ( 犃狉犪犫犻犱狅狆狊犻狊狋犺犪犾犻犪狀犪 ) ( 犜狉犻狋犻犮狌犿犪犲狊狋犻狏狌犿 ) ( 犗狉狔狕犪狊犪狋犻狏犪 ) 5 ( 犌犾狔犮犻狀犲犿犪狓 ) 5 ( 犅狉犪狊犻犮犪犮犪犿狆犲狊狋狉犻狊 ) L ; U [2] X%& ; H!! ' "#X ',!' *+ $ `abc% &' *Z[.\ 1 ; U! U * 31N; U, X 5 ; U < Marquez [4] * ; UH! ()*,+, 41N-. / 0,1 > 17 N ; U Y (1) ; U_.\ - 2 AtCYP38 L AtCYP20 3! 3 4 ) 5 Ⅱ (PSⅡ)X67 _8 9L : ; [. \ AtCYP38 R XT ;,_ < PS Ⅱ: => L?@, ^_8 9: X=>;,A 犃狋犮狔狆 38B( ' CD E [5] KX AtCYP20 3 R ;,! F2,G PPIase 8 9. \ X $ H R I J2 KL8 9 $_LMA ' NL `abco) XSN [6] 2 LMA 8 8 (2 CysPrxs),= > 8 L 2 P Y,AtCYP20 3, _2 CysPrx.\! QNSN 8 9 E X R [7] AtCYP18 3 AtC YP19 1L AtCYP19 3^_ - 2 犃狋犮 狔狆 18 3B( S T U L E U V 2 X W X.\>Y,! UL => X Z[L\ [8 9] 犃狋犆犢犘 18 3 犃狋犆犢犘 19 1L 犃狋犆 犢犘 19 3!]^ _`,A!ab 犃狋犆 犢犘 18 3Xc [9] ;! ' * + * Z [. \ 犃狋犮狔狆 71 B( \ ' *+ T Q ( (' Li [10] %&* At CYP71 _ H 3 U LO *',A ^_ UK; U 犃狋犮狔狆 40B( N % T * + L Q ( [11] Smith [12] %&* AtCYP40 _ HSP 90. \ X' *+ X AtCYP20 2^_ NADPH X;,R : QNX[ 2 P [13] H 2 BZR1 (brazzinazole resistant1,bzr1) U 2 X (,>< BZR1_ T U 犉犔犇 ( 犉犾狅狑犲狉犻狀犵犔狅 犮狌狊犇, 犉犔犇 ) 2 XC R $ 犉犔犇 T,A 犃狋犮狔狆 20 2B( T [14], X AtCYP21 3 L AtCYP21 4= > P Q X PPIase,! X, LIJ.\ [15] %&* ; ^_ ` L X ` a b c AtCYP19 2! L ` `a [16] ;AtCYP19 4 b L` `a [17] ; 犃狋犆犢犘 19 1L 犃狋犆犢犘 57^_ I 9 0 J X D b [18] ;AtCYP40 _ U microrna >,_ : X ; [19 20] AtCYP18 3 _ VirD2.\ IJ [21 22],A _0 J AvRpt2 X >,> < X [23] Li [24] % & * AtC YP b RPM1X RIN4, *IJ RPM1 V2X Db ; _ - 2 > AtCYP18 3^_ _ - 2, X' L*+ [21] ; X AtCYP20 3 ^_ -, _ I J K [25 26] ; K X AtCYP20 1, _.\^_' - 2L X [27], X AtCYP59L AtCYP65R T ;,AtCYP59 _ RNA Ⅱ.\! Lper mrna [.\, A AtCYP65 _ F a *. \ [28 29] AtCYP19 4 AtCYP20 1 AtCYP21 1 AtCYP21 2 L AtCYP21 3^_,G AtCYP19 4! K L,, $ U2^_ [30] 2 ; U U >29N; U,_ S H! ()*,+, 46N-. [31], XY %&, 1 7N UXY,

3 $%&78 Table1 Thereportedcyclophilinsandtheirfunctionsin 犃狉犪犫犻犱狅狆狊犻狊狋犺犪犾犻犪狀犪 U Gene U Genelocus IJ Subcelularlocalization UY Function ^ Reference 犃狋犆犢犘 18 3( 犚犗犆 1) At4g38740 Chloroplast _( VirD2 0 J AvRpt2 % RIN4.\;^ _ S T % E U X- 2 Interactingwith 犃. 狋狌犿犲犳犪狊犮犻犲狀狊 VirD2protein, 犘. 狊狔狉犻狀犵犪犲 AvRpt2proteaseandtheplantRIN4protein;in volvedinhormone(brassinosteroid)regulation [21 24] 犃狋犆犢犘 18 4( 犚犗犆 5) At4g34870 IJK Cytoplasm b `abc Respondingtolightstress [9] 犃狋犆犢犘 19 1( 犚犗犆 3) At2g16600 IJK Cytoplasm ^_ I 9 X Db; b ` a Involved inplantdefenseresponseagainst 犘. 狊狔狉犻狀犵犪犲 ;respondedto lightstress [9,23] 犃狋犆犢犘 19 2 At2g21130 IJK Cytoplasm! L ` ` a Inducedunderhighsaltstress andlowtemperaturestress [16] 犃狋犆犢犘 19 3( 犚犗犆 2) At2g56070 IJK Cytoplasm _IJ CaM. \; b ` a b c Interac tingwithintracelularcalciumproteincam;respondingtolight stress [49,9] 犃狋犆犢犘 19 4 At2g29960 K ER ; Golgiapparatus, $ U 2 GNOM ^ _ In volvedinvesicletrafickingbymodulatingthefunctionofgua ninenucleotideexchangefactorgnom [30] 犃狋犆犢犘 20 1( 犚犗犆 7) At5g58710 K ER ^_ X $ Involvedinproteinphosphatase2A (PP2A)regulation [27] 犃狋犆犢犘 20 2 At5g13120 Chloroplast ^_ NAD(P)H : QN; BZR1 X ( A ( T Involvedin NAD(P)H dehydrogenase complexformation;causingconformationalchangeinbrazzi nazole resistant(bzr1)protein (thereby alteringflowering patern) [13 14] 犃狋犆犢犘 20 3( 犚犗犆 4) At3g62030 Chloroplast I 8 9- a* LMA ' NLb Db,_ SAT1(LMA ' N )> ;^_ - Link inglightandredoxsignalstocysteinebiosynthesisandstress response,interactingwithsat1 (essentialenzymeincysteine biosynthesis);involvedinjasmonatesignaling [6,25 26] 犃狋犆犢犘 21 3 At2g47320 Vacuole Mitochondria ^_ X,! L I J. \ Involvedinthepermeabilitytransitionporeof mito chondriaandplayingacrucialroleinnecroticandapoptoticcel death [15] 犃狋犆犢犘 21 4 At3g66654 Chloroplast Mitochondria ^_ X,! L I J. \ Involvedinthepermeabilitytransitionporeof mito chondriaandplayingacrucialroleinnecroticandapoptoticcel death [15] 犃狋犆犢犘 38 At3g01480 Chloroplast ^_ )5Ⅱ : L 8: X; L ;PPI ase InvolvedinassemblyandmaintenanceofPSⅡ and oxygenevolvingcomplexes;ppiaseauto inhibition [5 7] 犃狋犆犢犘 40 At2g15790 IJK Cytoplasm ^_\ ' ; U,HSP90V2X RISC ;, Tombus : ; Involvedinvegetative shoot maturation;post translationalgenesilencing;hsp90 mediated RISC assembly; inhibition of Tombus virus replicaseassembly [11 12,19 20] 犃狋犆犢犘 57 At4g33060 IJ Nucleus 犃狋犆犢犘 59 At1g53720 IJ Nucleus ^_ 0 J X DbInvolvedinplantdefense responseagainst 犘. 狊狔狉犻狀犵犪犲, _ A (SR) L RNA Ⅱ X. \ ^ _ L pre mrna Involvedintranscriptionandpre mrnaprocessingviainteractionwithserine/arginine rich(sr) proteinsandrnapolymeraseⅡ [18] [28] 犃狋犆犢犘 65 At5g67530 IJ Nucleus _F a*.\ Interactingwithubiquitinligase [29] 犃狋犆犢犘 71 At3g44600 IJK Cytoplasm ^_ O * ', U, U K ; L H3 Involvedin organogenesis,epigenetic gene silencing, chromatinassemblyandhistoneh3 modification [10] _`abc (2) X OsCYP21 4 ^ _ L 8`abc % (ABA)- b Y +, PPIase % CsA X A,OsCYP21 4 = > PPIase 犗狊犆犢犘 21 4 % L H 2O 2 X,

4 4, :; UY %& 833 X> 2 >, R, $ 8 *Z.\ [32] OsCYP2. 8(ROS)!X $U2,! 0IJX8 P.\ [ X.\ Ruan [33] %& OsCYP2 % 8 X, ROS X! 犗狊 犆犢犘 2 `af2,!'" $ #, OsCYP2!G$`a. N ^_-,,] PEG % & ' & L ABA 犗狊犆犢犘 2 U 8 A K 8 _`, T[`a,! `a (L ( c [34] Kumari [35] %& *! 5 L 犗狊犆 犢犘 2 G % 8 ` a X Lee [36] X%& 犗狊犆犢犘 18 2! ( ()(* `a UX L pre mrna ) * ) ( L ` a. \ F 2 犗狊犆犢犘 18 2 L U 2 U 犗狊犛犓犐犘 X,OsSKIP _ OsCYP OsCYP18 2 I J K + I J ^ _ ` a UX, )(`ax 犗狊犆犢犘 19 4 T `af2,`,- 犗狊犆犢犘 c 10/% OsC YP19 CD E, 犗狊犆犢犘 19 4 X ', 0 1 L,, 犗狊犆犢犘 19 4 ' % 'cx [37] OsCYP19 42> PPIase, Lee [38] \ 3 4 L BiFC &567!` * G89 )*:' X OsCYP19 4ASE L 犃狋犚犆犖 1(+, 1 N X ) ; X 犗狊犆犢犘 20 2R=> PPIase X; U, L ` a U 5 L IJ `ax L `ax [39] Trive di %&* 犗狊犆犢犘 25 b T ` a b c,! ` L)( [40] 犗狊犆 犢犘 1 b `a.\, QNL ', A^_ ' *+ [41] V bc, ; ^_' -, Kang [42] %&* OsCYP2. 2 P^_' b X_,! (X < * Z [. \ 犗狊犮狔狆 2 B ( ' by,!( >T ' ' *+ 3 4 L pul down 7 = 6 OsC YP2 % _ > 2 P SKP1 X G2 2 OsSGT1, OsCYP2, _ OsSGT1 X.^_' - 2,%&* OsCYP19 4_' >, ` `a b U [37] 2 239:$%&78 Table2 Thereportedcyclophilinsandtheirfunctionsinrice U Gene U Genelocus IJ Subcelularlocalization UY Function ^ Reference 犗狊犆犢犘 1 LOC_Os01g02080 Chloroplast b `ab c, X ' Respondingtohigh saltstress,inhibitingtheformationofriceblastfungusgrowth [41] 犗狊犆犢犘 2 LOC_Os02g02890 IJK Cytoplasm ^_' -, (' ;, ROS?V ;! 5 L G T [`a Involvedinauxinregulatorysignalmechanism, regulateoflateralrootgrowth;increasingsalttoleranceofrice seedlingsbyrosremoval;increaseingthemultiplestresstol eranceoftobacco, 犈. 犮狅犾犻 and 犛. 犮犲狉犲狏犻狊犻犪犲 afteroverex pressed [33 35,42] 犗狊犆犢犘 18 2 LOC_Os08g44520 IJK Cytoplasm 犗狊犆犢犘 19 4 LOC_Os06g49470 ( )Lumen 犗狊犆犢犘 20 2 LOC_Os05g01270 Chloroplast ` a U X, ) ( ` a X Regulatingthetranscriptionofstress resistancegenes,andin creasingthetoleranceofplantstodroughtstress _' >, ' ` a b Involvedinauxin transportion,andsignificantlyrespondingtocoldstress ^_ L ` a Involvedinregulationof tobaccoand 犃狉犪犫犻犱狅狆狊犻狊 stresstolerance [36] [37 38] [39] 犗狊犆犢犘 21 4 LOC_Os07g29390 Mitochondria ^_, I J, b T ` a Involvedin regulationof mitochondrialpermeability,apoptosis,respon dingtoavarietyofstress [32] 犗狊犆犢犘 25 LOC_Os09g39780 ( )Lumen b `a Respondingtohightemperatureandhigh saltstress [40]

5 G$; TLP40R A XTY,^ _ '*'LIJ - 2. AtC YP38 B,H ) 5 Ⅱ E X, ^_ )Ⅱ TN W % K L TLP40 _ AtCYP38=>82%C D,AtCYP38 = > PPIase A TLP40 R Y PPIase, RU TLP40 >PQX PPIase L [43 44] _ OsCYP2 E X LeCYP1 I J ; <,^_ ' - 2, ' * + [45 46] 犔犲犮狔狆 1 B ( T ' E, bccd Y ( F_` 3G*+(E% 7 [47] Kaur [48] %&6; TaCYPA 1= > PPIase, 犜犪犆犢犘犃 1 5 %, G. \ - H? I TaCYPA 1 _ AtCYP19 3XC D 72.67%, %_ $ CaM *' 2JK.\ [49 50], ; TaCYPB ^ _ K X _ ;, X= ' _N [51] Yu [52] * ; TaCPY _ LM6VS/6AL N ) O ; RcCYP1 I J K = > PPIase,! KX. 2 P*Z[.\ [53] X CsCYP 1, _ TPR X8 9 U2_ RNA Ⅱ [54 55] X TaCYPA 1 [56] TX Catr1CYP [57] _ CsCYP 1=> X LC D, _ CsCYP 1 DXY,%&* TT X Catr1CYP [57] L P( 犅犲狋狌犾犪狏犲狉狉狌犮狅狊犪 )T X Betv7 [58] ; E9 < X ; ^ _ T ` a b c,q] ` )( -RL 9 StCyP X S T T ` a *,! ` )(L-R.\, -RL 0G mrna c A> U 9 X V [59] XTWX; LpCYP! -R ` L `a ST T,!Z[ X cy [60] ; ThCYP1 F G!IJ!IJ cet,hx c 38 L F2,! IJ E U [61] ; pcypb,^_ bl `abc,g % _W A Q N : X [62] 5 X C CyP>, 9 ' [63]! T'; AdC YP 9 ' X [64] Kinoshita [65] %& * VcCYP ^ _ I J X D b Z 犆犪犆狔犘 1! N X ( \ L T,A 9 F 2 [66] Kanakala [67] * ; NtCYPB = > X [ TYLCV \ X. \ TYLCV R ], LG$^._N`[ &,NtCYPB! ( TYLCV X A 2 TYLCV \, % & b. c, % ; ^_ - StCyP (MeJA)L ABA > b,! ABA. \ c [59]! 犌犼犆狔犘 1 * G (GA3) (IAA)L (ZT)- [68] ; U 犜犺犆犢犘 1 ABA > b [61] Mainali [69] 5; U < U,>> 62 N5; U( 犌犿 犆犢犘 1 犌犿犆犢犘 62),!18 * U > G,R / 0 E5X; U <, R S! U! X; U < %& 犌犿犆犢犘狊 G U E 'U,! LN >,A! * + X [70 71]!` a b c, L 5 ; mrna ST> E * ' 0 ` a, ; mrna A5; 2 >( `a, ; mrna ST X 0 Y, '`a = D [72] 3 XG$; U GY 4 ; =>T IJ LY,V ^_ KX N_. 2 P*Z PPI ase ^_ DNA _ %. 2 X,, ^_ X PL8 9 $ X' *+,!`abc%

6 4, :; UY %& ; 23"#$%&78 Table3 Otherreportedplantcyclophilinsandtheirfunctions ; Cyclophilin B Sourcespecies UY Function ^ Reference 犅犲狋狏 7 P 犅犲狋狌犾犪狏犲狉狉狌犮狅狊犪 S T E9 Apolenalergen [58] 犆犪狋狉 1 犆犢犘 T 犆犪狋犺犪狉犪狀狋犺狌狊狉狅狊犲狌狊 S T E9 Apolenalergen [57] 犜犔犘 犛狆犻狀犪犮犻犪狅犾犲狉犪犮犲犪 L. ^_ )5Ⅱ E X L T K, _ K > Involvedintheregulationofphoto system Ⅱ specificproteinphosphataseactivity,involvedintheregula tionofphotosyntheticsystem Ⅱinavarietyofkeyproteindephosphory lation,andplayingaroleinproteinprocessingandtransportion [43 44] 犔犲犆犢犘 1 犔狔犮狅狆犲狉狊犻犮狅狀犲狊犮狌犾犲狀狋狌犿 ^_ PSⅡX; L3% _ A.\ Involvedin assemblyandmaintenanceofpsⅡ andinteraction withphosphatasein thylakoidmembrane [45 47] 犜犪犆犢犘犃 1 犜狉犻狋犻犮狌犿犪犲狊狋犻狏狌犿 5 %,_ I J CaM. \,^ _ PPIase8 9 X Increasingtheheatresistanceof 犈. 犮狅犾犻,interacting withintracelularcalcium protein CaM,involvedinregulationofthe PPIaseoxidationandreductionmechanism [48 50] 犜犪犆犢犘犅 犜狉犻狋犻犮狌犿犪犲狊狋犻狏狌犿 ^_ X> _;,^_N Involvedintheeficient foldingandassemblyofproteinsubunits,involvedinwheatmaturation [51] 犜犪犆犢犘 犜狉犻狋犻犮狌犿犪犲狊狋犻狏狌犿 _ LM6VS/6ALN ) Associatedwithdiseasere sistanceofthewheat 犎犪狔狀犪犾犱犻犪狏犻犾狅狊犪 6VS/6ALtranslocationline [52] 犚犮犆犢犘 1 O 犚犻犮犻狀狌狊犮狅犿犿狌狀犻狊 L.! KX[. 2 P * Z [. \ Playingcriticalroles intheproteinrefoldingasmolecularchaperones [53] 犆狊犆犢犘 犆犻狋狉狌狊狊犻狀犲狀狊犻狊 _ X J b PthA Tdx % RNA Ⅱ. \ InteractingwiththebacterialefectorproteinPthA,theplantTdx, andrnapolymeraseⅡ [54 55] 犛狋犆狔犘 犛狅犾犪狀狌犿狋狌犫犲狉狅狊狌犿 b L 9 Respondingtostress,hormone, mechanicaldamageandpathogen [59] 犔狆犆犢犘 WX 犔狌狆犻狀狌狊狆狅犾狔狆犺狔犾狌狊 bt `a,! Respondingtomultiplestress,highly expressedinleaves [60] 犜犺犆犢犘 1 犜犺犲犾狌狀犵犻犲犾犪犺犪犾狅狆犺犻犾犪 Increasingthesalttoleranceoftobacco [61] 狆犆狔犘犅 犞犻犮犻犪犳犪犫犪 L. _ CsA X, L% F2 Inhibitingtheac tivityofcalcineurin byinteraction with CsA,induced bylightand heatshock [62] 犆 犆狔犘 5 犅狉犪狊狊犻犮犪狆犲犽犻狀犲狀狊犻狊 b',=>, 9 ' Respondingto biologicalstress,havingantifungalactivitytoinhibitthegrowthoffun galpathogens [63] 犃犱犆犢犘 T' 犃狉犪犮犺犻狊犺狔狆狅犵犪犲犪 E 9 ' X Increasedtheresistanceof transgenictobaccowithitsecotopicexpressionto 犘犺狔狋狅狆犺狋犺狅狉犪狆犪狉犪狊犻狋犻犮犪 [64] 犞犮犆犢犘 犞犻犮犻犪犳犪犫犪 L. ^_ IJX Db,!IJ9'K Y(L T Involvedinthedefensiveresponseof 犞犻犮犻犪犳犪犫犪 beancelsand much moreexpressedincelprotoplaststhaninrootsandleaves [65] 犆犪犆狔犘 1 Z 犆犪狆狊犻犮狌犿犪狀狀狌狌犿 L. b, 9 % XF2 Respondingtostress,induced bypathogenicbacteriaandplanthormones [66] 犖狋犆犢犘犅 犖犻犮狅狋犻犪狀犪狋犪犫犪犮狌犿 L. ( X [ TYLCV, TYLCV \ Changing thetranscriptionalpaternofyelowleafcurlvirustylcv,andreduc ingthetransmissionoftylcv [67] 犌犼犆狔犘 1 犌狉犻犳犳犻狋犺狊犻犪犼犪狆狅狀犻犮犪 ^_ 2 * ', b b c Involvedinseedgermination andplantgrowth,respondingtohormones [68] 犣犿犆犢犘 犣犲犪犿犪狔狊 L. b` ` a b c, mrna * ' S T Respondingtolowtem perature,highsaltstresswithmrnaaccumulation [72] 犌犿犆犢犘 5 犌犾狔犮犻狀犲犿犪狓! LN,!, b` `abc Ex pressedinyoungshootsand matureplants withhigherexpressionin youngshoots,respondingtolowtemperature,saltstress [69 71], &'*Z[.\ G ;. `abc ^_ X- 2,,Q] ; %_T ] : 8 L 8.\, ; ^_, 2 XIJ,A ; ^_ IJ FL(IJ

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