,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 "#$%#&' ()*+,-./01,MELVIN

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$@ A B C D E F G"#HIJK LM,NO( 犕犲犪犲狇狌犻犪犵犾狔狆 ) ( 犆狔狆犿犲犻犪犳犻 )2/ P7Q R ST,UVW(XYEF 7,Z2 P7 [\I]^,_$`ab c ; :(1)7U,[(1.73±0.21)kg m -1 s -1 MPa -1 ] A[(0.47±0.02)kg m -1 s -1 MPa -1 )], 745689[ 犘 50=(-4.$

1 ,2017,37(4): 犃犮犪犅. 犅犲犪犾. 犗犮犮狀. 犛犻狀. : (2017) 犱犻 : /j.issn "#$%#&' ()*+,-./01,MELVIN T.Tyree (./01 /, ) : :; 4 7 < & = >,% A B C D E F G"#HIJK LM,NO( 犕犲犪犲狇狌犻犪犵犾狔狆 ) ( 犆狔狆犿犲犻犪犳犻 )2/ P7Q R ST,UVW(XYEF 7,Z2 P7 [\I]^,_$`ab c ; :(1)7U,[(1.73±0.21)kg m -1 s -1 MPa -1 ] A[(0.47±0.02)kg m -1 s -1 MPa -1 )], [ 犘 50=(-4.99±0.11)MPa]V[ 犘 50=(-2.28±0.09)MPa)] (2)7E [ 犇 mean=(13.61±0.46) μ m)] [ 犔 mean= ( ±125.31) μ m] 9 [ 犇 h= (15.57±0.05) μ m]a[ 犇 mean=(9.48±0.18) μ m 犔 mean=( ±78.56) μ m 犇 h= (11.68± 0.02) μ m], E [(2260±77)N/mm 2 ] A [(2970±171)N/mm 2 ] (3) E Y 6 7 U (4)Ca 2+ 2 P 7 6 " # _, V E 6, $ 7$ a :E ;45689;; 2345:Q :A 犚犲犾犪犻狀犺犻狆犲狑犲犲狀犆犪狏犻犪犻狀犚犲犻犪狀犮犲犪狀犱犜犪犮犺犲犻犱犃狀犪犿狔牔犕犲犮犺犪狀犻犮犪犾犜犪犻犻狀犕犲犪犲狇狌犻犪犵犾狔狆犪狀犱犆狔狆犿犲犻犪犳犻 LIU Mengmeng,MELVIN T.Tyree (ColegeofForestry,NorthwestA&FUniversity,Yangling,Shaanxi712100,China) 犃犪犮 :Thecavitationresistanceofgymnosperms,anindicatorofdroughtresistance,dependsonthexy lemtracheidstructureandmechanicaltraits.inordertoprovidetheoreticalbasisforreasonablecultivation duringproduction,westudiedthewater usingstrategiesoftwogardenspecies 犕犲犪犲狇狌犻犪犵犾狔狆 and 犆狔狆犿犲犻犪犳犻 withtheadvancedcentrifugetechniqueandcomparedthetracheidanatomy andmechanicaltraits.theresultsdemonstratethat(1) 犕. 犵犾狔狆 hashigherspecifichydraulic conductivity[ 犓 s=(1.73±0.21)kg m -1 s -1 MPa -1 ]andweakercavitationresistance[ 犘 50=(-2.28 ±0.09)MPa]than 犆. 犳犻 [ 犓 s= (0.47±0.02)kg m -1 s -1 MPa -1 ; 犘 50 = (-4.99±0.11) MPa].(2) 犕. 犵犾狔狆 haswiderandlongertracheidwithhigherhydraulicdiameterbutlower density[ 犇 mean=(13.61±0.46) μ m; 犔 mean= ( ±125.31) μ m; 犇 h=(15.57±0.05) μ m;density= : ;: : (Z )!: (1991-),,!"#$,%&' $ $ "# E mail:liumengmeng1006@126.com (:MelvinT.Tyree,)*,+!$,-,%&' $ $ "# E mail:mtyree@163.com

2 4(, : EF (2260±77)N/mm 2 ]than 犆. 犳犻 [ 犇 mean=(9.48±0.18) μ m; 犔 mean=( ±78.56) μ m; 犇 h= (11.68±0.02) μ m;density= (2970±171)N/mm 2 ].(3)Thetorus margoof 犕. 犵犾狔狆 is moreflexiblethanthatof 犆. 犳犻.(4)Ca 2+ weakenedtheflexibilityoftorus margo.duetothenar row,shortandlessdensetracheidwithlessflexibletorus margo, 犆. 犳犻 islesscavitationresistant andmoretoleranttodroughtstress,comparedwith 犕. 犵犾狔狆. 犓犲狔狑犱 :tracheid;cavitationresistance;anatomy;flexibility J [ a [<&7 I,_:7<& 4F,745689A 4[\I]^XY I,50% 7 B C D( 犘 50) ; BC D, 2789 A 2 [1],%&?@AE F [2 3],E,, [3] (torus margo),sperry_6a [3] ;Pitermann [4] XY7U ;Jansen [5] P 6 a, 6 U R Bouche [6] 115 P P 7, U,6a 7, 2a@ , "#%&a [1,2,7], " # : a L 2 P 7 / P,%&$ 7!", #_ / $ % P L P & & ', & 4, BCDC7, (_b c) 1 *+ LM 1.1 :;,-./A./01010,. aad, 23 4,5 ' : ; MJ/m 2,7 2105h,712.9, (7 <)25.8, 3 (1 <)-1.1,7=> 42,7=> -19.4, 0? 309d, 10? 4185,@ A B 206d, C(221d,76;651mm,D7$ 6;E 7 6; _79% 1.2 :;?./01010$ 7 ( 犕犲犪犲狇狌犻犪犵犾狔狆 ) ( 犆狔狆犿犲犻犪犳犻 ), _ 20 7 H 7FG _2~37$FG 1.3 <,=> 1.3.1?"@$AB0CD HIJK L_5~8 mm 7FGMN OM 7 +Pa, QRV'7!SN E [ T,O U V W - X D P? 5 Y 2cm 7FG,I BCD, ( 犓 h), 犓 h= 犉 /(d 犘 /d 狓 ), a 犉 _ ;,d 犘 /d 狓 _ /BCD,EK7 9 犓 h Z*[\]?( 犃 s)7 U U _ ^ U, ( 犓 s M), ( 犓 s T)_ : Y Hagen Poi seuilel `ab7,c _: 狀犓 s = (π/128η ) 4 犇犻 / 犃 s (1) 犻 =1 a 犇犻 :E 7, η : 25 7 QRST\ I S 7 Cochard J K M [8 9] A 2 756, IM 56 [10 11] H LM K,D P? 5 Y6mm, 27.4cm 7 FGS JK _1000r/min ( A 9_ MPa) 7, _,( 犓 max),jk' 7 9( 犜,MPa), 9 FG7, ( 犓 hi),\ic `, [ ( 犘犔犆 ): 犘犔犆 =100 ( 犓 max - 犓 hi)/ 犓 max (2) QRST\I WeibulS : 犮 ( ) 犘犔犆 100 =1- 犜犈犡犘 - (3)

3 a 犮 :, 犜 _ 9 \I7S T`,12% 50% 88% 7BC D 犘 12 犘 50 犘 88 2 P F G E()*+1 < BCDE E 9 9 <:I (LeicaRM 2235;Leica,Nussloch,Germany) ;, [ ] K? _ 16μm 7,DP 5 I _ 0.1% S, NH a,i 7(30% 50% 75% 90%) S A 7,I 100X ( _ ImagerA2;Zeiss,_Infinity1 5C,Lume neracorporation),i WinCel2013a (Regents InstrumentsInc.,QuebecCity,Canada) ;E ]? ; D]K?[ 73 ],D] :TLBCD KL ;,D F G [ ] ; 1000~ 2000 E Y E [ ] ]? ( 犃 ) ( 犇犻 )7`bD E 7 犇犻 =2 犃 / 槡 π E (Density)= 犖 / 犃 m, a, 犖 _ ; ] 7,E, 犃 m : [ ] 7 ]? A E 7 ( 犇犻 ) MJ be 7 9, HIE 7 9 犇 h ; 9 `c : 1cm 7,, ; 犓 max, 9, 9( kPa);7, ( kPa),D 9,,7 ; `L M K,, 犓 r= 犓 h/ 犓 max _ # (10 mmol L -1 KCl! CaCl 2 ) 6 7," " # I2PFGI600kPaS 6h, D#0.2~0.3h ; 犓 h 6h $ % & L,9 _0; 犓 h,i, '7,XU; 2 P GH4I I ±> ;, EQR STXY7 P 7I StudentT Test S -, [ I MicrosoftExcel S 2 [ 2.1 "#$%#1J?"@ 1(,7^ U, U, ) A, ^,:7 2.3 *,,:73.6*,;V 犇犻犇 h = (4) 槡犖 A, 7LU,` D E7 9U<,U,7 犎 W = 4 犇犻犃 m (5) BCDE :IJ M [6] FG7E ( 犔 ),[? 5 Y _6mm, 1cm 7 F G,I [_W, N80% 30%% 1 1U 7 b a,60 a 48h,I E?1 J,,I WinCel2013aS ( 狌犿犪犵 )*+1F0!FG,,,,a, 7 % I E Y 6 7 FGHLM,,O FG a +,-; PY )( 犘 <0.05), +,- ; P,( 犓 s M) U,( 犓 s T) )( 犘 <0.05), 1 7^ U,( 犓 s M) U,( 犓 s T)7UV Diferentnormalletersindicatesignificantdiferencebetween species( 犘 <0.05),diferentcapitalletersindicatesignificant diferencebetween 犓 s Mand 犓 s Tofthesamespecies ( 犘 <0.05),thesameasthefolows Fig.1 Thecomparisonoftheoreticalandmeasured specifichydraulicconductivityof 犕. 犵犾狔狆 and 犆. 犳犻

$z{ijj«５３４μｍ [ １３１５μｍ Uij ]9 z Ã Ä Ç ` 8? s U X １８１９ μｍ Uij? ４ [ ç è ９ ¹ } { ¹ [ [ ij ３２１μｍ [ ９１０μｍ D i Ý ] 9 Ç78?sU X １３１４μｍ U ij [çè １２７ ÝW0 Òz{ }{U ２２ 456754RuvwxH z{ } { U ]? ２ s Ｗｂｕ \ µ!$ $O p q 0 D z Ó Ô １２ Ôâ U\Ø IX$ OP#$ T k l $ Û ÝWUòó ô6 G¹]^ í _ `ÈÝW ± â ^#%& IX T ５０８８ Õ UfghzØ 犘１２犘５０犘８８ [ 犘１２ PY²êQ7 ì : R S Õ U f g h Ü Ý 犘５０ ô6tu]zwvôõ Ufg h Ü Ý 犘８８ zw W j V Ô Õ U f g h Ü Ý$

4 -.. ß z{ }{RSTVW ij klmnop]u Ð z GÄH ２ #$; e IX e; øj) ²ij UKW ¹LM² UKW i j N ß U ( 7 U; １２３６８±０２２ｋｇｍ１ z{ ｓ１ＭＰａ k øu ２ B }{U; D z X １６３±０１６ｋｇｍ１ｓ１ＭＰａ１ k ø １ U ３ B Y }{ KW8 ez{ ７４７ z{ijj«５３４μｍ [ １３１５μｍ Uij ]9 z Ã Ä Ç ` 8? s U X １８１９ μｍ Uij? ４ [ ç è ９ ¹ } { ¹ [ [ ij ３２１μｍ [ ９１０μｍ D i Ý ] 9 Ç78?sU X １３１４μｍ U ij [çè １２７ ÝW0 Òz{ }{U ２２ RuvwxH z{ } { U ]? ２ s Ｗｂｕ \ µ! O p q 0 D z Ó Ô １２ Ôâ U\Ø IX$ OP#$ T k l $ Û ÝWUòó ô6 G¹]^ í _ `ÈÝW ± â ^#%& IX T ５０８８ Õ UfghzØ 犘１２犘５０犘８８ [ 犘１２ PY²êQ7 ì : R S Õ U f g h Ü Ý 犘５０ ô6tu]zwvôõ Ufg h Ü Ý 犘８８ zw W j V Ô Õ U f g h Ü Ý １３ z { U 犘１２犘５０犘８８ î １６１２２８３０５ ¹}{U î z { ± ＭＰａ犘１２犘５０犘８８１９６２７１３２７ＭＰａ犘１２犘５０犘８８ U X aôa Y １ Y z { Q R S T U V W Ëë}{ [z GÄUYj]±e}{ ２３ 45675IijkBCGy=EFGH z{ }{f g h i j k l Z 0 Ô A? ３ z{uij e } { i j ª? ２ z{ }{U ] ±e}{ [ij k } { U １４３ B i j Ｆ２Ｔｈｖｕｎａｂｖｓｏ犕犵犾ｇｙｕ狔狆 }{U １１７ B ijª }{U ７６ Y ２ａｎｄ犆犳犻 z １ 45=75Ic4d{ }~ t Ｔａｂ１Ｔｈｏｍｐａｓｏｎｏｎｓｏｎｏ犘犔犆ｏ犕犵犾ａｎｄ犆犳犻狔狆Ｓｐｓ ÓÔ １２ ÕfghÜÝ 犘１２ＭＰａ ÓÔ ５０ ÕfghÜÝ 犘５０ＭＰａ ÓÔ ８８ ÕfghÜÝ 犘８８ＭＰａ z{ 犕犵犾狔狆１６１±００６ａ２２８±００９ａ３０５±００７ａ }{ 犆犳犻３５７±０１７ｂ４９９±０１１ｂ６３２±０９２ｂ a ËÌ«CDEYÏ$ ÔA 犘００５ âì ＮｏＤｎｎｏｍａｓｎｄａｓａｎｄｎｂｗｎｓｐｓ犘００５ｈｓａｍａｓｈｏｏｗｓｇｎ? ３ z{ Ａ }{ Ｂ ij ;? b5äuh Ｆ３Ｔｈｏｓｓｓｏｎｏａｈｄｏ犕犵犾Ａａｎｄ犆犳犻Ｂｎａｈｐｈｇ狔狆

5 "# %#E*+/OJR Table2 Thecomparisonofcharacteristicvaluesofthexylemtracheidstructureof 犕. 犵犾狔狆 and 犆. 犳犻 P Species 犇 mean/ μ m 犔 mean/ μ m 9 犇 h/ μ m Density/(N/mm 2 ) 犕. 犵犾狔狆 13.61±0.46a ±125.31a 15.57±0.05a 2260±77b 犆. 犳犻 9.48±0.18b ±78.56b 11.68±0.02b 2970±171a 4 7E 97[? Fig.4 Thefrequencydistributionoftracheidineachbinclassandhydraulicweightof 犕. 犵犾狔狆 and 犆. 犳犻 ^T;9,@T;9 5 7,5 % Thesolidlinesmeantheprocessofincreasingpressureand thedashlinesmeantheprocessofdecreasingpressure Fig.5 Thedynamicchangeofrelativehydraulic conductivityof 犕. 犵犾狔狆 and 犆. 犳犻 /7 [3], E XY 6 7 a, 6 V 7 P,,7U6 V 7 P,,V,_ 7 6 AB ; 9, 7 67C 9,7, 6 I600kPa 6h7,5 % Fig.6 Thedynamicchangeof 犓 rofflushing 犕. 犵犾狔狆 and 犆. 犳犻 withtwosolutions under600kpafor6h VD, 7,VC,E 7 AB /; 9,7, ad,e 76 ABW/,; 76( 5) 2 P,I 10 mmol L -1 KCl,A CaCl 2

6 4(, : EF ,-; PY )( 犘 =0.05), +,- ; PI )( 犘 =0.05) 7 I600kPa 6h 7,(Recovery 犓 r)uv Diferentnormalletersindicatesignificantdiferencebetween species( 犘 =0.05),diferentcapitalletersindicatesignificant diferencebetweenflushingsolutionsforthesamespecies( 犘 =0.05) Fig.7 Thecomparisonofrecovery 犓 rafterflushing 犕. 犵犾狔狆 and 犆. 犳犻 withtwosolutions under600kpafor6h ( 6),; KCl 2PE Y6 7 Y 6h,' 7,& A, : Y, 2P7,, )( 6) L9 0,,& A( 7),; 67 I KCl 6h 9 _ 0,,[',746% 77%, I CaCl 2 6h 9 _0,, [ ', 7 78% 91%( 7),; CaCl "#$%#"4SLT@ MN0JR ""#;V 7 [/ 0 :G,]?,F ;, &V 7 [ / 0 G@$ H5, 7:I,]?,T 7 [,[/ 犘 50[ _ MPa, A :J P, : $ P, K45689 R, LJ 7$ ]^ [T,G@ K [/ 7;, $V 7 [/ 89 MN$ 7 & 45689,8OV,&J 8;,$[/ 89,$ C $ [ A, _ /$ % P, $ 7 $ a 3.2 "#$%#"4SLT@$MN0 ( )*+1 BCD /]?,E ;,E@ [/ 8 9,B C D 7, E,,7,, [0, R [14 15] 1 : A 7 B C D BCD [ 9,BCDE,, [ 9, B [ 0 ""#a,7e V, V, U,V, 犘 50; 7E V,V,U,V, 犘 50V 2 a, [BCDE a77 9%& D[, D[:E,%&@:E 7 ; D[7, 7 6K7 6 7 [2,7,12] " " # a, 9 EU & A 3.3 "#$%#U&' F ,,:, 32< 7,, [3],1! P 7 :6 /,'E SN7 E,P7E 56 Q 6, 7 9,!P ABE XY/R, P R ""#-S TU,U,6V,V R 6 7 ', 67,,W 6 K7 [4] 2a,FGaE 7 6 W "#,X*7 6O /,,Y*76/, a P [13],S7 # 2 P 7 ; b 7 KCl,, I CaCl 2,V ( 6) [E XY '$Z, :, 6: 7, : [ 7 C,7\] 7^,S

7 [ 9,Ca 2+ K + 8O 1 ^_, : Ca 2+ 8O C`7 ', ^ [16] YFG7BCD7, V, X^- P [16 17] Ca 2+ 2P B67, a ,1bS-S VW6: [1] CHOAT,B, 犲犪犾.Globalconvergenceinthevulnerabilityof foreststodrought[j]. 犖犪狌犲,2012,491(7426): [2] CAI,J,SZHANG,MTTYREE.Acomputationalalgorithm addressinghowvessellengthmightdependonvesseldiameter [J]. 犘犾犪狀犆犲犾牔犈狀狏犻狀犿犲狀,2010,33(7): [3] SPERRYJS,MT TYREE.Water stress inducedxylemem bolisminthreespeciesofconifers[j]. 犘犾犪狀犆犲犾牔犈狀狏犻狀犿犲狀,1990,20(13): [4] PITTERMANN J, 犲犪犾.Therelationshipsbetweenxylem safetyandhydrauliceficiencyinthe 犆狌狆犲犪犮犲犪犲 :theevolu tionofpitmembraneformandfunction[j]. 犘犾犪狀犘犺狔犻犾犵狔,2010,153(4): [5] JANSENS, 犲犪犾.Plasmodesmatalporesinthetorusofbor deredpitmembranesafectcavitationresistanceofconiferxy lem[j]. 犘犾犪狀犆犲犾牔犈狀狏犻狀犿犲狀,2012,35(6): [6] FRANKLIN GL.Preparationofthinsectionsofsyntheticres insandwood resincomposites,andanew maceratingmethod forwood[j]. 犖犪狌犲,1945,155(3924): [7] HACKE UG, 犲犪犾.The HydraulicArchitectureofConifers [M].SpringerInternationalPublishing,2015: [8] COCHARD H, 犲犪犾.Evaluationofanewcentrifugetechnique forrapidgenerationofxylemvulnerabilitycurves[j]. 犘犺狔犻犾犵犻犪犘犾犪狀犪狌犿,2005,124(4): [9] CAIJ, 犲犪犾.Whathappenswhenstemsareembolizedinacentri fuge?testingthecavitrontheory[j]. 犘犺狔犻犾犵狔犘犾犪狀,2010, 140(4): [10] LIY, 犲犪犾.Evaluationofcentrifugalmethodsformeasuring xylem cavitationinconifers,difuse andring porousangio sperms[j]. 犖犲狑犘犺狔犾犵犻,2008,177(2): [11] CORCUERAL, 犲犪犾.Phenotypicplasticityinmesicpopula tionsof 犘犻狀狌狆犻狀犪犲 improvesresistancetoxylemembol ism (P50)underseveredrought[J]. 犜犲犲,2011,25(6): [12] SPERRYJS,UG HACKE,JK WHEELER.Comparativea nalysisofend walresistivityinxylem conduits[j]. 犘犾犪狀犆犲犾牔犈狀狏犻狀犿犲狀,2005,28(4): [13] FICHOTR, 犲犪犾.Vulnerabilitytodrought inducedcavitati oninpoplar:synthesisandfutureopportunities[j]. 犘犾犪狀犆犲犾牔犈狀狏犻狀犿犲狀,2014,38(7): [14] FAN D,ZXIE.Severalcontroversialviewpointsinstudying thecavitationofxylemvessels[j]. 犃犮犪犘犺狔犲犮犾犵犻犮犪犛犻狀犻犮犪,2004,28(1): [15] SCHULTE PJ,AC GIBSON.Hydraulicconductanceand tracheidanatomyinsixspeciesofextantse[j]. 犚犲狏狌犲犆犪狀犪犱犻犲狀狀犲犇犲犅犪狀犻狇狌犲,1988,66(6): [16] ZWIENIECKIMA,PJ MELCHER,NM MICHELE HOL BROOK.Hydrogelcontrolofxylem hydraulicresistancein plants[j]. 犛犮犻犲狀犮犲,2001,291(5506): [17] NARDINIA, 犲犪犾.Ion mediatedenhancementofxylemhy draulicconductivityisnotalwayssuppressedbythepresence ofca 2+ inthesap.[j]. 犑狌狀犪犾犳犈狓狆犲犻犿犲狀犪犾犅犪狀狔, 2007,58(58): (:)

;54 ;1 ( ' ) Vol.54 No JournalofXiamenUniversity (NaturalScience) Jan.2015 doi: /j.issn RSTU V ; &! Y,,, [,2\

$;54 ;1 ( ' ) Vol.54 No JournalofXiamenUniversity (NaturalScience) Jan.2015 doi: /j.issn RSTU V ; &! Y,,, [,2\$ ;54 ;1 ( ' ) Vol.54 No.1 2015 1 JournalofXiamenUniversity (NaturalScience) Jan.2015 doi:10.6043/j.issn.0438 0479.2015.01.011 RSTU V ; &! Y,,, Z,@ [,2\] ( 8+ =,>^ E G,Y7,>^ 361102) ; : ( 犕狔犮犪犾犲狆犺狔犾狅狆犺犻犾犪