, ) (
*
:
:
E-mail: [email protected]
مﺪﻨﮔ ﺪﻳﺪﺟ و ﻢﻳﺪﻗ مﺎﻗرا ﺶﻨﻛاو ﺎﻫ
ي
،لﺪﺘﻌﻣ ﻖﻃﺎﻨﻣ
ﻊﺒﻨﻣ ترﺪﻗ ﺶﻫﺎﻛ ﻪﺑ ناﺮﻳا دﺮﺳ و مﺮﮔ
ﺮﻫﺎﻃ ﺪﻤﺤﻣ ﻦﺴﺤﻣ
1ي ﻠﻋ ، ﻲ ﺪﻤﺣا
2ي ﺘﺳﻮﭘ ﻢﻇﺎﻛ و* ﻴﻨ
3ﻲ
! "# $% #% &# '(
)
*+ , :
- / . / 01 2 '3 , 4
: 5 / 1 / 00 (
هﺪﻴﻜﭼ
/ ! "# $
%&'
' ()*
+, %-.0 ($1 .
$3
$3 $&.! $- 4
$5 6$7 (
4 8 6 9 .
%3 :;1) ' < 1
$=- ;> ?> %@0 +@7 (3 A.0 : 4 + 7 6 9 6$7 %=
3 B3 <C- D! ' 7 ;1$> 3 .
@9 0 3 ;$'E
% $#@9 !
$ :F> 7 , $G? ('$?> H? $$ A.0 &' 9 %&' IJ K LMIN 6$5 : .
$1$O ' P*>$Q %>
R $ 6 9 +7 )
4 6$7
69 * : D!$ ' 6 9 + 7 (
R B3 <C- )
@- :
7$
*Q VW
%1 @9 ' X ' (
A! $ 3 > :3 Y@3
$1 H $3 %Q$ P$> (
4 . Y$Z 3! [\- 3 B3 <C- ] :%@> ! 3 $=3 ($E ^ '
% 3
?#3 ,3 Y$Z _E (* B3 ;1$> ($1$O .
' 7 ;1$> $3
1 :` & :"**Q & B3 <C- D!
' ( B3 PQ@> ( $
Q$ ;"Q 1$ 3 = +a .
^@b c
CO2
' ( Q XH&
"**
;1$> B3 <C- ;"Q $3 3 Q$
$? >
d*> ] 1
% 1 3 "#
$\0 * '
( 3 "**Q 3 ]e %@Q"
. H@O& ;>
\de
$1 E (
@- ' B3 $ *C9 %7$3 3 $=H B3 <C- $O 3 = :f@*# 6$7 *Q 7$
] :$9 ( 3 3 3
% H@O& ( ( .
($1$O $O& $
;1$>
% 1$ $3 =$g H@O&
) B3 <C- 3 (
% .
]
( 3 B3 <C-
% ] "1 ' ( Hd$- :3
% (
.
ژاو ه يﺎﻫ يﺪﻴﻠﻛ : B3 <C- :
83 K
"#
: Y$Z 3! 0 :
'$9 $ (
+a B3
:
\de H@O&
E ($1 :
$ 6 9 .
ﻪﻣﺪﻘﻣ
! "#$% & "#'#() *+$ & ,) ' & -./ # #'01
( 2 3
4*!
3/
. ) (
! # (
*+0 /
& 67 8 301 * (
#$%
. 9;) -!
FAO (2008)
#/ "* <
=
% (& ?1 &
-! "#$% ,) 301* & #'
8 < @#A+4 4
(8 #!
B #$ "((
/ % @#A+4 4 -! ,) #$% (8 ?1 &
,) *D E/+ -1 3/ "F *G#0 "*
% < *!#! ! "F #$% *D (G#( & *+1
3 8 & 67/ ,) *D H ;I #'
01 #J(JK
L#!
.
#!
3 *
#' &
"
? & #(!
(0M! N$! -; 67/ *D 1 K .
# 3 *
#'
#O P#$ -! &
N$! +(
I#1 *D Q7
L#!
. '#)F # F -1 -+D &
-./ L ,
#' J #! #A+O
K % ;+/+I
L ) SN 3 K (
;+/+I -1 3(I*U -(/! # 3I#
1 )
";M 3 K (
V$
L#!
(Cruz Aguado et al., 1999)
-% WX
.+ "#JJK
3/ ! .
Y -!
-1
Simmons et
al.
(1982)
*! Z*! [\ #'#( *]
VO
Z*! [\ -1 9;) *1 -.8#7 ,) "& <#0I *) -*
-
#' Y -!
.
H'#1 ' .
^ -D(8#
-+' Z*! [\ ,#
& _`
H'#1 a#! ,#O & D #$ -*
. "&
-
#' *) . I*Y &
4*! <
(JJK
(Slafer & Savin, 1994; Richards, 1996;
Borras et al., 2004)
.! ";M & -1 #! *!
*D 1 *+1 +1#I *+$ &' ') &
3/ - . -+N8 7(K ? <VO HJ -7!
E! #D+/ #'#( -1 #() L & -*
SN # ";M
L -1 +' $ ? <
L -+I*) # #N )
Zhu et al., 2004
.(
Zhu et al.
(2004)
-1 9;) ,) *! 0 #&F SN H'#1 ? -* Z*! [\ b *Y &
E/
-^`
&
*]
. -+L - *D *!
*4 SN H'#1 3/
-^`
&
+I -O#/
H'#1 a#!
. L *D .
-1 *G -.8#7
<3/ -+I*) cd ,) % V O ,#O *!
- "& *+0(! 3(/# e #+
- *D -^(+
V O ,#O -! 3N % ,#O "#0
'
)
Kruk & Calderini, 1977
.(
,) - *D ?(#+` *((f
&
- . *((f b *Y )
67/
( *' "& #
F 3/! - )
Cartelle et al., 2006
.(
G+N'
*D (! L (! - . -
g(h
#' 3/ F 3/! ,)
(Slafer et al.,
1990; Fischer, 2008; Cartelle et al., 2006;
.
Calderini et al., 2006)
.
! #! 3J(J (
#+ *+0 e
,) - *D -1 i h8 ;(I c#.8#7 -! j!*
<3/ K ";M k# ') & _`
3J!#7
. "*N% 3 -! - . H ;I <% #!
-! *^ k#(J+ -0(' - "& H'#1 b *Y & "F *D ?(#+` H ;I
*)
(Slafer et al., 1996;
Slafer, 2003)
.
Slafer et al.
(1996, 2003)
! (
"#
H'#1 - "& <- . H ;I #! -1
l(AM #$ - - . H ;I -1 <!#
a#! -D! < H'#1 - *' -! ;+/+I -m) "& ?(#+` H'#1 3O
#' -N/ #$+
( 3/ L ; .
*! -+I*) cd c#.8#7 <# *' -!
bY# *! L nod ,#O SN 3 K p+M
(Ma et al., 1990; Koshkin & Tararina, 2003)
"#0 e #+
' -#*! Y -1 od #'
. 3$%
-
#' 3 K <,) *+0(!
3/ -+I# H ;I SN .
Fischer & HilleRis (1978)
% V O -+/ -! ,#O -1 #G' ;(
*)
! F 3/! $!#0 e #+ *1 .
< *!#!
#! ,#O *! SN 3 K -% ((. 3$%
& + #! c#+ *D ?(#+`
"#D
#'
"#&
#' 3I*) *$! +M
"
i e #+
(! # *G -J7 -! -J7 i & + -! H #&F V(. ,#O .
' "1#
( c#.8#7 ;( "* <-7!
L ,#^ K
(Esmaiel Pur, 2007; Jan
Mohamadi, 2006; Ahmadi & Judi, 2007)
*J -1 k#N
H'#1 , -! #$F -' .
#N -! *D
& . [\ b *Y & SN cO H'#1 #'#(
Z*!
#' #L ,) SN 3 K N -^(+
3/ L .
m' ( #J+ H ;I <c#.8#7
+K #` <L *(4q *` ^ r#! (L4 S0.0 ;+/+I H ;I <Z*! ?(I*1 7 *L (2 ,) *D s ? r -% &
3/ L 9;) .
NO L ,#^ c#J(JK <# *' -!
*+1 "#D "F -1 -; E *L G' "*
-! -0 [*Y E(K 3!Y b(O #'#( t
3/ -+I*) cd <-+L % Z*! [\
. \8
"#'#() 301 <u#4 3!Y *+1 3$% *X# -.8#7 L ,#^ ) cd -!
. c#.8#7 <*G [*Y &
-+I*) cd NO !F (8 *` VO i & #$ "*
& *v1 # VO
) VO i *' -! !F VO i
V ( /*! *Y#4 (' -! <3/ L #+/
?(#+` #! W+ ,#O & c#+ (8 #'
O ( V
#' 3/ L Q#M+ 01 p+M .
*!#!
*D N 3(/# /*! *X# -.8#7 & ['
#+` #! ,) V O % ,#O - (
8 ? ( <c#+
01 ,*) +. <*/ bY# *! L nod [\ b *Y & SN cO H'#1 #'#( -! H1
! Z*!
.
شور و داﻮﻣ
ﺎﻫ
& ?AI *X# H't`
w B -YK
 D0 c##N nod 3& *) -#M) L *% x*1 SO "*$ #G0 .
0 #&F *'
-(1 W# -! D(+/o`
w *7O B +#/
*+
W# # -1 <!
B +#/
-; u#4 & *+
) +(/
/ ( _y/ L *`
B +#/
*+
"F #OI
z(* -! (/` 1 u#4 "#' & YM #!
^ 3N -!
B -!
L*`
. i(+/o` *' -
& { -! #' |/
L -(N. 01 .
{ -!
") -! 3U#
H!# ;( -#+& "NM & #'
-0 *O "L i ; (L4 V(J+
-! #'
co` (! *(! #}I <-; E *L u#4 #! #'
W# # H1 B
+#/
*+
L (L`
. ' ,#G
#F *! ,&r #((L 1 * #J <301 *+! &#/
L 0 #&F *' -! c#N/#K ~#/
. ?4 0 #&F *'
B d -O#/ #! i *' ,) -!
-^` -(J! L -+L\) O#!
& +I -O#/ -* #'
L [\ -OY ?K .
"#& (' o -!
p
#' &*' L -+L*! "#( & +/ (% #! ;(
.
!F +I -O#/ -* & ?NO # (
# cd -! AK
+I -O#/ -* & .! # <L ,#^ #*
) (8
Z*! [\ -*
( # L *+1 cd -! QF <
L 0 #&F *' -! & 3(I*U -! "(/
.
m' ( )% { -! <
(
* MN &
( u#4 67/ & *
") YM #! #$F 67/ <#'
#! L &
! -/# <
( #
?O *7O -! H1 ; B
/ +#/
L` *+
( L .
,) VO 3' & ?D0+ ? +1#I N(1* #'#(
n*Y - #` *! -1 ! Z*! [\ cL 3' "#
u!
#' { *D -/ #! I#A ?#1 (
*) V .
<\! nod -/ co HM! #0 #! /*!
O -/ & ,#O (
I*U #! "* d V (
*D 3
L Q#M+ c#+
. <( ,#O <{ (' -!
-! -.L L <*/ bY# V O ,#O & z(*
N Y -! <,*) +.
#` *D #!
( ,#O *
# *$L bY# % ,#O & z(* -! "*2 &#+0(` <
#+` #! <,*) +. <*/
( 8 ? ( *' -! <*r#!
Q#M+ (8 V1 V VO i " -! */ VO L . 3(N8# N -% ((. { -! <(m'
cL <";M -! SN cO Z*! [\ p+M #'
L ,#^ -*
)
% .(
-* W*L "#&
+I -O#/
) -*
~#(J
Zadoks et al.
(1974)
(
L ,#^ Z*! [\ -* (8 .
! #1 9
#` -1 ! cd (
Z*! * z *! L [\ #'
"#'#() <#( # cL
<
i L )*!
+I*) *{ '#L " -! M 3/ #(
L - .
Z*! ?#1 [\ -1 Z*! [\ -* ( *! <! #'
B % #'#( &
<
-* "# #` )*!
- *(L -* &#F #!
) -*
=
= &
_1& ~#(J (
*J -1
*K! #! [#A k#N
*`
3I*) cd <3/ - "L .
W^ -1
67/ 3' <-*
?(D0 Z*! [\ cL
.
,$' 1 2
"
34 567 894 $& )
!
"# $% &'( )*
1L
+
!
&'( ,-% "# $% &'( )*
+.
/!0 ,(
1L-nL
1 " 0 /$
+ )
&'(
+ 4 /56(
(
"# $%
2L
8 " 0 /$
+ )
&'(
+ 4 /56(
( 9:4 ,-% "# $%
+.
/!0 ,(
2L-nL
; " 0 /$
1 )
&'(
+ + 4 /56(
(
"# $%
3L
<
" 0 /$
1 )
&'(
+ + 4 /56(
( 9:4 ,-% "# $%
+.
/!0 ,(
3L-nL
= ! ) (
!
"#
#` { -!
f K H ((
% QF c*
I*`
( ?
-* & <L ?AI Y u#4 ' -N/ ?
-!
8+ Y B
I#A Y -! #(!F *' & ?NO #!
& <
b B +#/
*+
) -0 -./ b (
-
*! u#4 -(/! ;^ ") -/ u#4 & #( *'
*)F #G+/
8F [*U 3L*!1
( ( *O
L .
*'
& -*
) (
* *{ *D i " -!
L -+I*) .
-D(P#^F & < *!#!
B -*
&
*()
?#L *{ & ;( #( *' <L ,#^ 3!Y B
*D
*) .
&
-
#' 3!
D ` ( *O & H
"F +I*) )
°C
c -! <
3#/
( D
cd "F & _`
& d _y/ <3I*)
3!Y
-
#'
& *I bNY
*) -N/#K * :
100 ﻚﺸﺧ كﺎﺧ نزو × -
بﻮﻃﺮﻣ كﺎﺧ نزو
%=
ﻧزو ﺖﺑﻮﻃر ﻲ
ﻚﺸﺧ كﺎﺧ نزو
$% &' ()
V^` <,#$2 #' -+' V2*` Z*! ?(I*1 +I -O#/ & .! V0L )
[\ #'#( # "#&
Z*!
( +/ & #+/ #!
?(I*1 #G +/ *+
(SPAD-502, Minolta, Japan)
&
*() L . !
*' {
-/ ?(I*1 +K Q#M+ -!
#$F V2*` Z*! r#! #( <( #` 3O (G#( 3I*) *O /*!
" -!
& #(.
L -+I*) *{ V2*` Z*! ?(I*1 .
1. Auger
*
$% &' +,
&#) cr#N *! Z*! [\ *] /*! 3$%
) 3 ' <;+/+I -&
(...
O#! V2*` Z*!
<#
B #G+/ & #+/ #! Z*! [\ & _` &
IRGA
Z*! % #( '#L #'#( LCA4
V2*`
&
*() 3I*) cd 1\ c#d .
!
& _` { 3O #G+/ "*1 *N(8#1 V({
-{K ?4 V2*` Z*! 7/
&
*() )
*N2 (
#.I & lM0 K i L *O
;+/+I )3
B
*! S!* *+ *! *D(
-(#]
( & .!
B -! *! -Y!* -(#]
(G#( 3N] c*1 *' I#A
*() *) .
- ! ./(0
*!
&
*() -! ?1 <c#d
#' ,)
0 #&F *' % )
B *' -!
#! i -O#/
d ( "L i04 & _` 3L*!
-
#' &#/%
-
#' { -! 3I*) cd 3/ -(/! -N/ &
;' "& *D ((.
L & - .
(G#(
- .
L F*! ;( -N/
.
!1
,* & #+/ #! F 3/ -! e #+
#F ;I
#'#( (G#( (m' <L _# - ;^ SPSS
- 2 "&F & #+/ #!
#F 67/ D
B / L - #J .
V/ *!
,* & ;( #' ;I
Excel
L #+/
.
2. Infera Red Gas Analyzer
3. Photosynthetically active radiation
ﺚﺤﺑ و ﺞﻳﺎﺘﻧ
!
"#
#J ,#^
-1 *1 lM0 D 9 -! c#
( *! Z*! [\ # (
I*` % QF ";
( u#4 ?
;I *] '#L -! 3N 0
3L )
?DL .(
*+0(!
#( -! j!* u#4 3!Y s ";(
1L
!nL
* #/ Z*! [\ "! '#L #( #! *{ & -1 c# Z*! [\ #( n7/
. +L . #! \8
Z*! & */ i [\
& H(! ' J & #'
E/ u#4 3!Y -(M QF Z*!
#' *. "# *%
3/ L -+/#1 .
3/ -(X*I #! '#' e #+
*L -1 b *Y & "+! #L +!Y 3 K E
[\
Z*!
#' # ( - "L*` &#! V1
QF 3I ' *. ";( u#4 3!Y ' ' S!# % -I*d "#D b *Y & H'#1 V'*I K !F .
;( *G S!#
)
Richards, 1993; Zhu et al., 2004
( -! Z*! [\
SO ) z/# "#&
( -! `#1 #+4#/ V({ *!
3!Y 3 K "#& 0 L H'#1 { -^(+ QF [*A #1 H ;I #'#D' & D L 9;) *+0(! *D 3L*! l4#L A 3/
. 67/ D(+h #D+/ S!# (m'
N$! { -! Z*!
3K *D QF [*A #1
3/ L #$0(` +!Y 3 K E *L
.
(P=0.01 <%> ) <=> ?@ >
2 C
34 567 )
2 D+ 34 567nL
"
4
,$' 2 9 E
( FG H4# " -! 4 .
$
<=> $&> -9
;
%2 34 567 2 " 4
-9 H9 "I J!K F-L 57 " "
&.
$&-?>
. 2! /(0
/(0
*! Z*! [\ VO ?!#J+ *] /*!
*D c#d & i (' -8
#' "F . N
)
% .(
O % V O ,#O *G c#N -!
V(
#'
$!#0 #+I -.8#7 #+/ p+M 4 & SN cO H'#1 p+M n7/ -! H1 "#0 .
*{ -! *!#!
3(N8# -% -1 /
SN -{o ?!#O c# ,#O ";M -+L
L#!
. F 3/! e #+ (' ;( '*) c# #J
c#
. 0 ,#O #+I (!
) "#0 e #+
3/ 0 .(
,$' M 1
>
4 LI "-+N C O> P2 Q-
> P > % RST
$&
!"
#$%
&'*
+-* ./ 0! &12 345ns
/
378 9/9* :/9ns 7 &*
**
/
: /**:/:** ; (A)<=
8>>**
/
3?4 ;5/5>5ns 99/** ; (B)@&$ ABC 7DDns
/ D?
7>/;8Dns E;/ns A×B 88?
/
8> E8/DEE >/DE3 4; FG
*
: **
.
: ns
!
c#
#' . (!
E *L ,#O *D
H(! ' % ,#O *D L '#0 0 #&F ! V O ,#O &
)
?DL .(
% -! ,#O H1
"#0 SN cO H'#1
' . V1 ,#O (!
(8 ) O ( * L V' "F L VO ;% -!
& i (' -! *D H1 <Z*! [\ #(
n7/
0 SN cO H'#1 .
*D H'#1 E/+
L#
O ,#O Z*! [\ &
B
%
% ,#O
% *(8*` ,#O ! )
% (
~K *D H1 SN cO H'#1 -! *
"#0 .
*{ -! *!#!
( #` *D -1 /
#+` #! ,#O 3 K 3 -! ( #` *D ?(
-% ";M 3 K #! kr#+ L#N SN .
-!
(8 V1 ,#O #! - #J (8*` ,#O -2*) <#*' -+L *+0(! H1 EJI ;( ,#O + 8 <
U
cL -!
/#` SN cO H'#1 r#! #' / "F *G#(! -^(+
;( (8*` ,#O + -1 3
3/ -% ";M 3 K #! - *D
(Kruk &
Calderini, 1977; Reynolds et al., 2005 & 2007)
.
H't` < *!#!
-! #'
*! t ?(#+` H ;I
& kr#+ ";M 67/ % ? -! -% <(8 8
*r#! 3 SN 67/ % ? -! 3N
3/ 4*!
.
#'*+#` -% & -;' "& .
I*. z(* -! -1 +' ";M #! EN*
& &
";M
"F 3(8#.I &
L#!
.
.$& ) > % "<=> 1 D%
9
<=> <4 RS2 - F-L 57 V D) $7 +
$>
&.
H?>
.
H+
) % R W2 $T
$&
34 567 $ 2 4 HX?> 34 567 Y-Z "2
* IJKL M
NO%*
$ P%Q R1LS T= &12 &'* + +/ $ UVW @&$ ABC
+/ &12
) 0+1X &/ T&X
( T=
3L
3L-nL 2L 2L-nL 1L 1L-nL 9
/
+/ / +:/ / /9 / +/ (&J WF%<V+=)
/
/ / +/9 /9: :/ :/* V&N / (&J WF%+V+) :
/
+/9 +/: +/9 / / /* / (Z+L- WF%<V+=)
/
+/ +:/ 9/ / /* /* !LO[/ (Z+L- WF%+V+) :
/
+/ +/ +/ / / / / (T&X WF%<V+=)1- /: / +/ / /99 :/* 0&\/: (T&X WF%+V+)
9 /
:/ / /9 / / / / (<V+=<V)&J
/
/ / +/ /* /9* /* /9 Q : *
"# $% &
) '(%
* /
* ) .(P=
, -0! 12
"
3 .(
! -
c#Io+4 .
- . *{ & ,#O (!
F 3/ -! - ;' "&
)
% .(
/*!
G+N' .
-N/ - . (!
3L % - ;' "&
) ww / .(r=
-! */
<( #` - *D ?(#+` #! V VO i "
3L - ;' "& *+0(!
. . *{ & VO
. "& (! _D -7! ! *+1 - L ;( ,#O * #/ (! - )
?DL .(
z7
! - -(/
Acreche & Slafer (2006)
D L (
G+N' -(% *! 8#+ ? r &
(! 3!#O % -
#'
*` Q\%
L *1q
3/
. ,#^ i h8 ;(I c#.8#7 <% #!
SN #D+/ -(& L
#! ";M
3J!#7 .
<=> <4 (P=[/[; <%> ) 4 R?@1 D%
"
P > $.2 >
$& Y-Z )
$ &LO$ ]^K* &$
K -&
.
) (
)(
) (
*' ~*+/ *` H ;I <#v " -!
-mN/ & . [\ b *Y & - <-N/ i #'
-mN/ - "& *(]
3L # O#! #'
(Ma et al., 1990; Cruz Aguado et al., 1999;
.
Cartelle et al., 2006)
. [\ <;( *X# H #&F
*] Z*!
. 3L - "& *!
)
% .(
# SN H'#1 + <*G c#N -!
%
Z*! [\ , -*
) & ( ?#1 Z*! [\
#'#(
<
<i -* )*!
& _` &
') ( - "L*` *K! -1 <L #
3/ -+L ` - "& H1 .
< *!#!
(!
-
#' % /K 3!#O *` Q\%
-+L -
#' & W#NL ko#1 E *L ,)
L *`
1 )
Borras et al., 2004
.(
& i (' *! Z*! [\ VO ?!#J+ *]
c#d
"&
- . .
0 )
% .(
-!
V(O -! b.+ ;( % V O ,#O *G c#N #'
c#d *{ & -.8#7 #+/ p+M cO H'#1 p+M n7/ -! H1 $!#0 #+I "#0 4 & SN .
67/ - . *! Z*! [\ #( *]
#+
B / . -! #$ H1 -+N8 <!
Z*! [\ r#! cL )
1L 1L-nL
( L j!*
*] Z*! [\ n7/ * #/
. - .
+L )
?DL .(
*r#! G+N' - .
)
r=0.64
( - "& -! 3N )
r=0.35
( 3L *D #!
.
<=> <4 (P=[/[; <%> ) 4 R?@1D%
"
34 567 R$ Y-Z E^9 4 *&+ > $.2 ) ,$' 2 E
.(
-9 F-L 57 " "
"I J!K
"
RS2
&.
$&-?>
. S!#
*G ( ' -! ; (
! 3 ( 3N - . *+0
#L *D ((. - "& -!
-* L
((. *K! -* - . ((. i h8I 3/ L *1q *D
(Borras et al., 2004; Fischer,
2008)
. Z*! [\ #'#( </*! -D #!
-*
) +I -O#/ -*
( "#& k#N *J
*] # <L # - . "L 3!#] ?(D0 1 3L - . .
"
-1 3I*) -^(+
z* -! SN cO H'#1 & - . H'#1 3/ ! *+1 .
-1 & "#' -! <*G c#N -!
- . <L # Z*! [\ b *Y & SN H'#1 H'#1 , ? r & D kr#+ 3/ 0 *] + . *D Z*! [\ +1#I n7/ 4*! *]
#
2L
3/ !3L
.
$% &' +,*
*! Z*! [\ VO ?!#J+ *] (m' VO *]
V2*` Z*! &#) cr#N c#d & i (' .
0 . V O ,#O -(1 *G c#N -!
%
0 c#d ";( *{ & H #&F /*!
-!#
H'#1 #'#( -! #D H1 (m' <!
"#0 4 & SN cO .
*{ -!
;+/+I 3(I*U -1 / Z*!
#' -!
'#L "#'#() -1 3/ J "F & H(! z*
'#0 L ;+/+I 3(I*U & #+/ *v1
Z*!
! #' - ? F
.
?DL -1 7#' B
"#0
3*/ H ;I a#! Z*! [\ <3/ L ;+/+I Z*!
#' '#L #! - #J # O#!
#( H ;I -1 L Z*! [\ "!
*{ &2L
#F .
! . ;+/+I 3*/ 1\ #(
% [\ *] # 3I# H ;I '#L -! 3N
Z*!
1 "*N% #' .
3*/ H ;I #+/
3*/ <Z*! [\ #'#( -! H1 ;+/+I 3 ' *.
-&
4 & $!#0 H ;I ;(
3{ -1 3(8# < "#0
CO2
"
-&
#( t -!
H'#1 '#L #! - #J 2L
3I#
. ;+/+I 3*/ (! -!#0 H1 (2
3 ' -&
3/ L 9;) ;( *G S!# -1
(Koch, 1996; Richards, 1996; Cruz Aguado,
.
1999)
F*I -& V(J+ *+1 *! 38r *! ";M &#( ;( ;+/+I
-& "L &#! ";(
#'
3$%
CO2
. Z*! 67/ H'#1 *G [*Y &
;M &#( H ;I -8; -!
Z*! *! "
# O#! #'
3/ ";M cO H ;I #.
. & D <";M cO
SN ;+/+I 3*/ *! \)*(] ?
L#!
. -2*'
*1 Q\% *+0(! #/ <L#! *+0(! ";M cO '4 H ;I ";M SN (! #/ 3{ z(L 3(8#.I H ;I 3 -! k#+ #$ 3I#
(1 V ;F
SPS1
(` H ;I SN ;+/+I 3*/
1
(Reynolds et
al., 2005)
. 1 Y -!
"
*(] #! ";M -1 3)
3 ' -&
c#d <
-&
*+1
1
*(] #!
";(
CO2
"
*! ";M -1 3)
(I; 3 '
*+1 *F 3/ *] ;(
1 .
$% &' () 3
#$ V2*` Z*! ?(I*1 +K *! Z*! [\ *]
V0L -+' )
L # #` ?*
( # & _`
SN H'#1 #'#(
. L )
?DL
.(
7#'
'#0 -1
L L # #` ?*
) V0L -+' (
<-+I# H ;I SN cO H'#1 cL O -2 *' ,
3/ -+I# H ;I ;( V2*` Z*! ?(I*1 .
-D Y -!
#( -! j!* ?(I*1 +K *+0(!
& ! 1L
c# Z*! [\ n7/ * #/ '#L #( #! *{
. 3L . V$ ? & D -D -! -% #!
";( -^(+ Z*! *(` 3*/ *! \) *(]
(I*1 SN -7! #. <"F ?
";M L#!
(Rajcan & Tollenaar, 1999)
+I*) *{ #!
,) ";M 3 K 3(N8#
(Borras et al., 2004;
Fischer, 2008)
i [\ -1 / *{ -! (2
(! Q7 #. #^ -! *^ ;+/+I S!# */
3$% ";M SN (
! &#
( M *+0 L SN -! ";
3/ #+I *(4 -! Z*! *(` -^(+
. < *! o
+K *! 3Nv *] Z*! [\ L -+) -1 7#' 3L u#4 3!Y .
+K Q7 3(.X < *!#!
Z*! * #/ V2*` Z*! QF N O#! #'
#
+K #` *! kr#+ ;( L Z*! [\ "#'#()
1. Sucrose phosphate synthase
1 8#.I H'#1 #$F ?(I*
(
;F 3 V
#' I*1 ( *(] &o
3/ -+L\) 3Nv .
1; D%
H $ _.2 H9 P-&9-N H9 <=>
>
"
H`a
CO2
>
"
(bc 34
$&
34 567 "2 )
,$' 2 E
( +
$ > <-N )9 7 $>
"
Oc -S
HN RT I .
H9 %2 9 <=> -9 @
-9 RS2 F-L 57 4 "
&.
$> $>
.
D%
U 1 (bc 34 DN+ R W2 $>
$&
Cd2 H!2
-S 34 567 2 , $.4 (L (&c eb "
34 567 "2 )
^K* &$
$ &LO$ ]
K `K .(
13 4()
*! Z*! [\ VO (! !#J+ *] #{+ [o4*!
0 "F ;% *D
c# #J '*) c# ;( L ,#^
. % ,#O H1
01 p+M bY# -! b.+ (m' ,) V O 0 '#0 .
[\
n7/ * L #$ Z*!
H'#1 . "#0 '#L #! - #J - *D
. -(/! SN cO H'#1 & *G c#N -!
! *+1 z* -! ";M cO H'#1 <Z*! [\
"#0 3/
' ,#O + <- *D -1
kr#+ ;( r#! (8 ?(#+` #!
";M 3 K #!
3/ -%
. H'#1 #'#( L -{o -1 7#'
<u#4 E/ 3!Y $G ";( N$! a#! SN #! - #J V2*` Z*! ?(I*1 #` ;+/+I W^ 3 #$ -1 < *) Z*! [\ "! #(
c*((f & *1 ? *I#! " -! ? L
*1 *()% 7 *L (2 *D
3/
.
REFERENCES
1. Acreche, M. & Slafer, G. A. (2006). Grain weight response to increases in number of grains in wheat in a Mediterranean area. Field Crops Research, 98, 52–59.
2. Ahmadi, A. & Beyker, D.A. (2000). Stomatal and non stomatal limitation of photosynthesis under water stress condition in wheat. Journal of Agricultural Science, 31(4), 813-825. (In Farsi).
3. Ahmadi, A. & Judi, M. (2007). Effects of timing and defoliation intensity on growth, yield and gas exchange rate of wheat grown under well-watered and drought conditions. Pakistan Journal of Biology Science, 10(21), 3794-3800.
4. Austin, R. B., Bingham, J., Blackwell, R. D., Evans, L.T., Ford, M. A., Morgan, C. I. & Taylor, M.
(1980). Genetic improvements in winter wheat yields since 1900 and associated physiological changes.
Journal of Agricultural Science, 94, 675-689.
5. Borras, L., Slafer, G. A. & Otegui, M. E. (2004). Seed dry weight response to source–sink manipulations in wheat, maize and soybean: a quantitative reappraisal. Field Crops Research, 86, 131–146.
6. Calderini, D. F., Reynolds, M. P. & Slafer, G. A. (2006). Source-sink effects on grain weight of bread wheat, durum wheat, and triticale at different locations. Australian Journal of Agricultural Research, 57, 227-233.
7. Cartelle J., Pedro, A., Savin, R. & Slafer, G. A. (2006). Grain weight responses to post-anthesis spikelet- trimming in old and modern wheat under Mediterranean conditions. European Journal of Agronomy, 25, 365–371.
8. Cruz-Aguado, J. A., Reyes, F. Rodes, R., Perez I. and Dorado, M. (1999). Effect of source-to-sink ratio on partitioning of dry Matter and 14C-photoassimilates in wheat during grain filling. Annuals of Botany.
83, 655-665.
9. Esmaiel Pur, M. (2007). Response of tow wheat cultivars to source size modification: interaction of cultivars and plant density under water stress and non stress condition. M. Sc. thesis. College of Agriculture, University of Tehran, Karaj. (In Farsi).
10. FAOSTAT. (2008). Agricultural Data. Food and Agricultural Organization of the United Nations, Rome.
Online at. http://faostat.fao.org/.
11. Fischer, R. A. (2008). The importance of grain or kernel number in wheat: A reply to Sinclair and Jamieson. Field Crops Research, 105, 15–21.
12. Fischer, R. A. & HilleRis, L. D. (1978). Effect of environment and cultivar on source limitation to grain weight in wheat. Australian Journal of Agricultural Research, 29, 433–458.
13. Jan Mohamadi, M. (2006). Effect of defoliation on yield and dry matter remobilization under various moisture and nitrogen regimes in bread wheat. M. Sc. thesis. College of Agriculture, University of Tehran, Karaj. (In Farsi).
[
14. Koch, K. E. (1996). Carbohydrate-modulated gene expression in plants. Annual Review of Plant Physiology & Plant Molecular Biology, 47, 509-540.
15. Koshkin, E. I. & Tararina, V. V. (2003). Yield and source/sink relations of spring wheat cultivars.
Field Crops Research, 22, 297-306.
16. Kruk, B. C. & Calderini, D. F. (1977). Grain weight in wheat cultivars released from 1919 to 1920 as affected by post-anthesis defoliation. Journal of Agricultural Science, 128, 273-281.
17. Ma, Y. Z., MacKown, C. T. & Van Sanford, D. A. (1990). Sink manipulation in wheat: Compensatory changes in kernel size. Crop Science, 30, 1099-1105.
18. Rajcan, I. & Tollenaar, M. (1999). Source: sink ratio senescence in maize: I. Dry matter accumulation and partitioning during grain filling. Field Crop Research, 60, 245-253.
19. Reynolds, M. P., Pellegrineschi A. & Skovmand, B. (2005). Sink-limitation to yield and biomass: a summary of some investigations in spring wheat. Annual Appl Biology, 146, 39-49.
20. Reynolds, M., Calderini, D., Condon, A. & Vargas, M. (2007). Association of source/sink traits with yield, biomass and radiation use efficiency among random sister lines from three wheat crosses in a high-yield environment. Journal of Agricultural Science, 145, 3–16.
21. Richards, R. A. (1993). Manipulation of leaf-area and its effect on grain-yield in droughted wheat.
Australian Journal of Agricultural Research, 34, 23-31.
22. Richards, R. A. (1996). Increasing the yield potential of wheat: manipulating source and sinks. pp: 134- 149. In: M.P. Reynolds, S. Rajaram & A. McNab. (eds.), “Increasing yield potential wheat: Breaking the Barriers’ Proc. Mexico, DF, CIMMYT.
23. Simmons, S. R., Crookston, R. C. & Kurle, J. (1982). Growth of spring wheat kernels as influenced by reduced kernel number per spike and defoliation. Crop Science, 22, 983-988.
24. Slafer, G. A. & Savin, R. (1994). Source-sink relationships and grain mass at different positions within the spike in wheat. Field Crops Research, 37, 39-49.
25. Slafer, G.A. 2003. Genetic basis of yield as viewed from a crop physiologist’s perspective. Annual Appl Biology, 142, 117–128.
26. Slafer, G. A., Andrade, F. H. & Satorre, E. H. (1990). Genetic improvement effects on pre-anthesis physiological attributes related to wheat grain yield. Fields Crops Research, 23, 255-263.
27. Slafer, G. A., Calderini, D. F. & Miralles, D. J. (1996). Yield components and compensation in wheat:
opportunities for further increasing yield potential. P.101-133. In M.P. Reynolds, S. Rajaram & A.
McNab, (Eds.). Increasing yield potential in wheat: breaking the barriers. Mexico, DF, CIMMYT.
28. Zadoks, J. C., Chang, T. T. & Konzak, C. F. (1974). A decimal code for the growth stages of cereals.
Weed Research, 14, 415–421.
29. Zhu, G. X., Midmore, D. J., Radford, B. J. & Yule, D. F. (2004). Effect of timing of defoliation on wheat (Triticum aestivum) in central Queensland 1. Crop response and yield. Field Crops Research, 88, 211–
226.
