免费文献传递   相关文献

Effects of Shading on the Leave Anatomical Structure and Photosynthetic Characteristics of White Clover

遮阴对白三叶叶片解剖结构和光合特性的影响



全 文 :!23" !3#
 Vol.23  No.3
! " # $
ACTA AGRESTIA SINICA
   2015$  5%
  May.  2015
犱狅犻:10.11733/j.issn.10070435.2015.03.031
wxÁÄ!::A.ymà~–ÝÐoÔÕ
ÕOT
1,
U*÷
2,
V+W
2,
M X2,œ%3
(1.0„®W-„à.ñí-../,ÿ0 ~3712100;2.0„®W-„à.×&-„./,ÿ0 ~3712100;
3.0„®W-„à.–¯./,ÿ0 ~3712100)
67

_×Åa]¡ùú8Ò=2
(犜狉犻犳狅犾犻狌狉狀狉犲狆犲狀狊)‰

 ‰50%j ‰70%NOÒ=22.ݙ€›j=HñI—\。€a+›:"#ia\ ‰SeIù
ú

Ò=22Ž

†+1+Íte

Ü«¶teùú

|‘Ý™€›z{v†4

õ=ûæ
(犘狀)ƒ;†+1
te

Ü«¶te¶犘狀 nœ‹‹¬··ù,¶|ýz{nœ‹l¬··ù。!› ‰£Ò=22.€›4—,
=û恃

89:

ia–

Ò=2

 ‰

ݙ€›

=JG
;<,-=
:S541.2    >?@AB:A     >CD=:10070435(2015)03065304
犈犳犳犲犮狋狊狅犳犛犺犪犱犻狀犵狅狀狋犺犲犔犲犪狏犲犃狀犪狋狅犿犻犮犪犾犛狋狉狌犮狋狌狉犲犪狀犱犘犺狅狋狅狊狔狀狋犺犲狋犻犮
犆犺犪狉犪犮狋犲狉犻狊狋犻犮狊狅犳犠犺犻狋犲犆犾狅狏犲狉
YANGWenquan1,CHUJipeng2,KOUJiancun2,ZHAOJiao2,HANMingyu3
(1.ColegeofLifeSciences,NorthwestA&FUniversity,Yangling,ShaanxiProvince712100,China;
2.ColegeofAnimalScienceandTechnology,NorthwestA&FUniversity,Yangling,ShaanxiProvince712100,China;
3.ColegeofHorticulture,NorthwestA&FUniversity,Yangling,ShaanxiProvince712100,China)
犃犫狊狋狉犪犮狋:Inordertolearntheadaptabilitymechanismofwhitecloverinthesystemoforchardandgrass,
theanatomicalstructuresandphotosyntheticcharacteristicsofwhitecloverleavewerestudiedunderno
shading,50%shadingand70%shading,respectively.Theresultsshowedthattheleafarea,upperepi
dermisthicknessandspongytissuethicknessofwhitecloverincreased,whereastheotherindexesofana
tomicalstructureand犘狀decreasedastheincreasesofshadingfromappletree.Theupperepidermisthick
nessandthespongytissuethicknesswerenegativelycorrelatedto犘狀,whereasotherindexeswereposi
tivelycorrelatedto犘狀.Inconclusion,theshadingchangedtheanatomicalstructuresofwhitecloverleave
andcausedthedecreaseofphotosyntheticrate.
犓犲狔狑狅狉犱狊:Appleorchard;Whiteclover;Shading;Anatomicalstructure;Photosyntheticcharacteristics
EFGH
:20140504;IJGH:20141013
KLMN
:¸
¹_ö-.2Ê¿¥
(31302027);¸ ¹iaÍ÷„…2ù¿¥(CARS28);ÿ0(-„—~à¿¥(2011KTZB0202);,Ú
®÷Í÷„…2ùՋ߿ÀÊ
(CARS3540);0„®W-„à.ÊË-̨×2Ê¿¥(2013BSJJ070)ÀÁ
OPQR

~Ó£
(1975),Å,ÿ0Ò¸§,ÊË,âÔ,mVÏО&j„ñ&øFñ֎IÜ.j-ÌRx,Email:wenquan05@sina.
com;:ex×Authorforcorrespondence,Email:hanmy@nwsuaf.edu.cn
  ͸˜Øن!¥àiañ͸,Z"Žj
Í"J*äØÙ²Žj²Í"I41.09%j
42.88%,|8,cYÑû4@˜Í¸ia4ào•
Í@8`žŽzàIÍ@
,2008$,Š@iaZ
"ŽjÍ"J*äø²Žj²Í"I48.0%
j38.9%,n_cYÑû4@®÷íîIºWÍ
÷
[1]。


Ýފ4@a–YBk.[\Ž~

ñ
òNO‹ÿ…,
9à4@O‰iaÍ"I¦Ñ¶
»ÀIo[2]。
èx$gïIa–`]´µvP
ƒa–YBڒ

4YB&5ñ=

‡5"k

†
‡#×/e

,Én?

¦Ña“Í"j»À

˜c
YÑû4@ia–ËÌjv)°’“I®ga–Ô
5´µ
[12]。
Ò=2
(犜狉犻犳狅犾犻狌狉狀狉犲狆犲狀狊)Ýޞ6SZ,J
ïñ2

âÂ2I{ÉéÓx¤

ŒŽ

|ˆùSh

­Ó‰¶a\&Ék

]|˜ˆbc+Œx¤o
•›œ

úgñ;#·

©­ñº
、^
añ/oÝ

n_‰cYÑûia–z©ªI]`

éfŠ@i
! " # $ !23"
a–àŽ$º`ž
[3]。
¥á

×
、`
][13],


fÒ=2jia\nMnIa]¡ùú8

Ò=
2·#ËÞia\I ‰NO

=à‹Ü

áñ·
Â2


·Þ
 ‰<Ò=2ÕÖIÌÍ<@Þ]4ùú6´v 
䒔

Ê8f§f]48|ý*] ¡<Ò=2
MòÕn

ñ·ûæ

Q«&À«n֎IÕÖ

æ<ÞW]€ùú8=à(<Ò=2ݙ€›
IÕÖÏdª¸|
[45]。
-×ÅÒ=2fa–8<
 ‰I©­ý5<Þ-.4Ô5ia–

Ñ>£¤
Ò=2j()a]¡ùúITˆñðâ~V 
¨

_N

?íî<ËÞia\Wh‹Œ ‰NO
IÒ=22.ݙ€›j=Hñ5WÌÍ

Ñ
ÅÒ=2
_a]`ž´
µI­¤$º¦R-.LH

1 STUVW
1.1 Ö×+Z[
íî4SÞÿ0~3¸Lx-„–-,Ú®
÷஖Iiaíî24

24w525m,$v²
àŽG2150h,$uvxy12~14℃,9øzÑx
y39~40℃,9øzƒxy-15~-21℃,$uv
É"621.6mm,КyUx34xÿ。íî4
YB_XÔY

1.2 Ö×kl
ia–ÕÞ2009$¡,a\W«4m,6«2m,
»`_

·82U”(Nagafu2),Wh`Ò=2,f"4
g·m-2,³e_1m,Þ2009$3%@f`。
íîPñ·fia–èèÿ ‰IÒ=2_<
à
(CK),¤=òØ TM201Lø)àe¼Þ9∶00,
12∶00j16∶00fa\Whñ·Ò=2I4Ö-R
 ‰Ý

ö°†G²

W‹ŒŽhKISOèI
=/

“uvð

x_ ‰Ë5I=à/e

Ë5Ⅰ:
`]Wè8hIÒ=2

=à/eÉ_<àI50%,
c50%I ‰Ë5;Ë5Ⅱ:` ]Wèè"èiaIÒ=2

=à/eÉ_<àI30%,c70%I ‰
Ë5

1.3 m®e@UVW
1.3.1 ÝY¨Èíî¢d 2013$6%,¤2Ž
âˆ2Ž

¤Žÿ–zâƒIÖ³
[6],
ˆ
xNJe

1.3.2 ÝájZYË FAA +ˆ,O—+ñ<
ð

šÎÔÕK.

†2.te

†+1te

Ö
DjÜ«¶te

ÖD«¶+Í·eQ³e

’
µ¼j2.«¶€›žJe
(structureclosely
degrees,CTR)jQne(structureloosedegrees,
SR):犆犜犚(%)=(ÖD«¶te/ 2.te)×
100%,犛犚(%)=(Ü«¶te/ 2.te)×
100%。
1.3.3 ÙIÛÜ 2013$6%,pר©ÿ¸IÈ
x

†ª10∶00—11∶00Ž,¤LI6400ܬµ=
â

ˆ‹Œ ¡NOIÒ=2=ûæ
(netpho
tosynthesisrate,Pn)、ÚFûæ(transpirationrate,
Tr)、ÍhCO2‡e(intercelularCO2concentration,
Ci)jxNŸe(stomatalconductance,Gs)。
1.4 ijkl
íîGH¤ Excel2003 5WfzË5,¤
SPSS12.0ÉO5WonewayANOVAÖãJK。
2 mnU,c
2.1 zn‘wxÁÄ!::vwU’ì)oÔÕ
Ý+1vÀ,"# ‰SeIùú,Ò=22Ž
ùú

+1IxNJeœ…

<à¶Ë5
hãRœ‹
(犘<0.05),-2 ‰Ë5hvÿœ‹
ãR

2.2 zn‘wxÁÄ!::A.ymàoÔÕ
Ý+1vÀ,ia–8`žIÒ=2I2.t
e

ÖD«¶te

ÖD+ÍI·e

ÖD+ͳ
e

ÖD+Í·³t

ÖD«¶Ü«¶·³tv
"#a\ ‰/eùúæ—4

<à¶2A ‰
Ë5h

ü50% ‰IÒ=2ÖD«¶+ͳe
¶<àãR‹œ‹C

|‘)z{ãRv}9œ
‹Éu
(犘<0.01);æ2A‹Œ ‰Ë5IÒ=
2

üÖD+Í·³tÿœ‹ãRC

2.te

ÖD«¶te

ÖD+ÍI·e

ÖD+ͳe

ÖD«¶Ü«¶·³tvîf9œ‹ãR
(犘<
0.01)。
"# ‰/eIùú

†+1te

Ü«¶t
e

2.€›Qneœ…ùú

2.€›žJe
†4

456
!3# ~Ó£: ‰<Ò=22.ݙ€›j=HñIÕÖ
q1 zn‘wxÁÄ!::vw1:A.ymàoÔÕ
Table1 Theeffectsofshadingfromappletreeontheleafareaandanatomicalstructureofwhiteclover
<àCK Ë5ⅠTreatmentⅠ Ë5Ⅱ TreatmentⅡ
2ŽLeafarea/cm2 1.0962±0.2880b 1.3760±0.3218a 1.4476±0.2501a
xNJeStomadensity/mm-2 524.400±70.3Aa 404.700±55.8Bb  368.700±36.5Bb 
2.teLeafthickness/cm 152.300±6.6Aa 129.100±4.8Bb 118.300±2.8Cc
†+1teUpperepidermisthickness/μm 17.920±2.09Bb 19.170±2.49ABb 20.330±2.25Aa
Ü«¶teThicknessofspongytissue/μm 51.750±4.41Cc 59.080±3.50Bb 68.080±3.13Aa
ÖD«¶teThicknessofpalisadetissue/μm 69.250±3.89Aa 39.670±4.44Bb 18.780±2.14Cc
ÖD«¶+Í·eLengthofpalisadecel/μm 31.670±3.43Aa 27.330±2.07Bb 17.950±2.19Cc
ÖD«¶+ͳe Widthofpalisadecel/μm 14.750±1.78Aa 14.330±1.46Aa 10.250±1.52Bb
ÖD«¶+Í·³tLengthandwidthratioofpalisadecel 2.160±0.24Aa 1.920±0.22Bb 1.780±0.28Bc
ÖD«¶Ü«¶tetThicknessratioofpalisadetissueandspongytissue 1.348±0.149Aa 0.676±0.106Bb 0.277±0.041Cc
2.«¶€›žJe犆犜犚 45.500±1.8Aa 30.700±2.9Bb 15.900±1.6Cc
2.«¶€›Qne犛犚 34.000±2.3Cc 45.800±3.1Bb 57.600±2.9Aa
  •:à4…wxJ*+Å犘<0.01j犘<0.05ÉuãR。Œ
Note:Differentcapitalandsmallettersindicatesignificantdifferenceat犘<0.01and犘<0.05levels,respectively.Thesameasbelow
2.3 zn‘wxÁÄ!:~–ÝÐoÔÕ
Ý+2vÀ," ‰/eIùú,犆犻,犌狊,犜狉 v
ß>Ñ¢ƒI”•

æ犘狀 œ…ƒ。50%j70%
 ‰ I Ò = 2 =  û æ J * _ 20.1,17.2
μmol·m
-2·s-1,S<àÒ=2ƒ‰21.7%j
33.3%。
q2 zn‘wxÁÄ!:~–ÝÐoÔÕ
Table2 Theeffectsofshadingfromappletreeonthephotosyntheticcharacteristicsofwhiteclover
<àCK Ë5Ⅰ TreatmentⅠ Ë5Ⅱ TreatmentⅡ
õ=ûæ犘狀/μmolCO2·m-2·s-1 25.74±3.04Aa 20.16±1.00Bb 17.16±1.73Cc 
ÍhÃp\će犆犻/μmolCO2·m-2·s-1 291.00±28.45Bb 325.60±13.65Aa 296.60±20.80ABab
xNŸe犌狊/molH2O·m-2·s-1 0.7914±0.2572b 1.3501±0.4511a 0.4506±0.1396c 
ÚFûæ犜狉/mmolH2O·m-2·s-1 11.156±2.814ab 13.858±3.248a  7.714±2.613b
2.4 Ä!:~–0NU.ymào80
¬·JK+›

+3),犘狀 ¶2.te、ÖD«¶t
e

ÖD«¶+Í·³t

ÖDÜ«¶·³tœ‹
l¬·
(犘<0.05);æj†+1tejÜ«¶t
enœ‹‹¬·
(犘<0.05)。
q3 Ä!:~–0N犘狀 :A.ymàod80i
Table3 Correlationcoefficientsbetween犘狀andleafanatomicalstructureofwhiteclover
õ=ûæ犘狀 õ=ûæ犘狀
=/Lightintensity 0.998 ÖD«¶+Í·eLengthofpalisadecel 0.928
xNJeStomadensity 0.992 ÖD«¶+ͳe Widthofpalisadecel 0.820
2.teLeafthickness 0.999
ÖDÜ«¶·³t
Thicknessratioofpalisadetissueandspongytissue
1.000
†+1teUpperepidermisthickness -0.988 ÖD«¶+Í·³tLengthandwidthratioofpalisadecel 0.999
Ü«¶teThicknessofspongytissue -0.973 2.«¶€›Qne犆犜犚 0.958
ÖD«¶teThicknessofpalisadetissue 0.997
  •: +Å犘<0.05Éu†Iœ‹¬·ñ
Note:significantcorrelationatthe0.05level
3 œ
3.1 zn‘wxÁÄ!::AmàoÔÕ
ž&2.<(NOI—\™š(ƒ

vP:
;_PID,4—:©­‹ŒI(

2.<(
—\I©­YZ2M

2+ŽHIj2.åUݙ
HII—。
f‹ŒI=à/e

_䖔

ž&
I2.€›ð⋌I©­ñ
[6]。
f/=

2.
+,_Šñ2HI

2.St

xNJeSà

ÖD
ǦSt

Ü«¶S:

æfÆ=

2.+,_
‰ñ2HI

2.S:

†+1xNJeS4

ÖD
«¶9‹’}

Ü«¶’}
[7]。
a2?¥Š

fia\ ‰¢

Ò=2I2.jÖD
556
! " # $ !23"
ǦS:

ÖD«¶+ÍSH

S4

ÖD«¶+Í
·³tjÖD«¶¶Ü«¶·³tS4

†+1
jÜ«¶St
,CTRS4,SRSà,2.«¶%
æSQn

!›Ò=22.ðâS/Ivîñ

PN
:©­‹ŒI=à(
[5]。
xN˜ž&]O2.ÉJ5qj5Wx2õä
I:|

0OÕ֞&IÚFx¤

xND,˜©­
(IýOg¥

xNIJeý(=à=GIÕ
Övà

"=/ùú

xNJeœ…ùà
[8]。
?íî
€a+›

"#=/Iùú

Ò=2+1xNJe
ùú

<`—\â£Þ4fSÑ=/ÝÞðâ
SàI2h+Žæ¾n< CO2 œ±I³.[5]。
-êšÌ
[5]
f ŠP8IÌÍ+›

Ò=2IxN
Jeçâf=/_10%j12%Ž,uS<àœ‹
ƒ

æ?íîfia–8IÌÍá’,

f50%
€=/Ž

xNJeۜ‹ƒÞ<à

!› ‰
ƒxNJeIŒŽ

#+›fv]´vNOj_öNO

 ‰
3.2 zn‘wxÁÄ!:~–ÝÐoÔÕ
"# ‰SeIù/

2ñ¥¯"ùú

=x
¤
。Gregoriou[9]<àá(犗犾犲犪犲狌狉狅狆犪犲犪) ‰2$¢Ì͒,, ‰30%,60%,90%¢,à
á\2. 犘狀 S<àƒ21%,35%,67%;–&

[10]
<Ò=2

XL\]
(犕犲犱犻犮犪犵狅狊犪狋犻狏犪)、]l
å
(犕犲犾犻犾狅狋狌狊狊狌犪狏犲狅犾犲狀狊)、,€l(犃狊狋狉犪犵犪犾狌狊犪犱
狊狌狉犵犲狀狊)5W40%,59%,77%I ‰Ë5¢,’,
犘狀 " ‰Iùú恃。=2=ûæ"#ia\ ‰SeIùú恃

50%j70% ‰Ë5J*t<àƒ‰21.7%j
33.3%。-˜,"# ‰SeIùú,‹Œž&`a
犆犻,犌狊j犜狉ߋŒI—\”•,5IŽh·H

ž&I`aJK¬·

–&
[9]
<
‹Œ*]5W40%,59%,77%I ‰Ë530d¢
’,

<ÞÒ=2

 ‰77%IË5犌狊 ,æ 
‰59%,77%IË5犆犻 ÷Ñ, ‰40%,59%IË
5犜狉 ÷Ñ,栉77%IË5°;-˜, ‰
<]lå犌狊,犆犻,犜狉 vÿœ‹ÕÖ。?Ó<Ò=2
IÌ̀a+›

"ia\ ‰SeIù/

Ò=2
犆犻,犌狊j犜狉 vß,q>÷Ñ¢ƒI”•。!›
ia\ ‰<Ò=2=ûæIÕÖýxNü¥j
™xNü¥IŒx¤

4 m
ÝÞia\I ‰

£ia–Wh`žIÒ=
2I2Ž

†+1+Íte

Ü«¶teùú

xNJe

2.te

ÖD«¶te†4

ÖD«¶
+͗H

—4

ÖD«¶+Í·³tjÖD«¶
Ü«¶tet†4
,犆犜犚†4,犛犚ùú;õ=û
恃

!›Ò=2v:;2.€›4—:©­i
a\I ‰

- ‰£|=û恃

žŸ>?
[1] öÕ÷,~Ó£,µ›[,.͸a–ñ]ÌÍ5“[J].]÷
-.
,2010,27(7):154159
[2] §mü,3Œ¦,–|ï,.hxÒ=2ùúYBol2Qol2į"IÕÖ
[J].]4.¸,2013,
21(3),485493
[3] ~Ó£,öÕ÷,µ›[.Wh`‹Œ]¢<„Åia\=
HñIÕÖ
[J].]4.¸,2011,19(1):2025
[4] HeidrunH,LarsW.Shadeavoidanceintheclonalherb犜狉犻
犳狅犾犻狌犿犳狉犪犵犻犳犲狉狌犿:Afieldstudywithexperimentalymanip
ulatedvegetationheight[J].PlantEcology,2011,130(1):53
62
[5] êšÌ,L›ñ.Ò=22.ݙvîñQ|<=/IÖ­
[J].]4.¸,2006,14(4):301305
[6] ×µ%,3/,·òó,.|Ñ# ‰NO3A‹Œ¢ëb
XL\]»`2.ݙ€›¶|=ñò©­ñ
[J].ž&ñò

,2012,36(4):333345
[7] JamesSA,Bel DT.Influenceoflightavailabilityonleaf
structureandgrowthoftwo犈狌犮犪犾狔狆狋狌狊犵犾狅犫狌犾狌狊ssp.犵犾狅犫狌
犾狌狊provenances[J].Treephysiology,2000,20(15):10071018
[8] L¯Ã,Nš,¹¬ó.þ`vîñ<‹Œ=à/eIÖ­
[J].­¤ñò.¸,2004,15
(2):201204
[9] GregoriouK,PontikisK,Vemmos.Effectsofreducedirradi
anceonleafmorphology,photosyntheticcapacity,andfruit
yieldinolive(犗犾犲犪犲狌狉狅狆犪犲犪L.)[J].Photosynthetica,2007,
45(2):172181
[10]–&,cÏ),¦™,. ‰<4`;-*]=HñIÕ
Ö
[J].]÷-.,2013,30(1):4451

%&( , -)
656