Effect of nutrient enrichment on growth and photosynthesis of the

! \)4A:;(4Bd(49C; KQQQ

O04C9 !((8D
C:+ C 8@4*6(4 +('4(CD( 08 XW =CD 03D(41(+H !C*(D 08 )60*0DE:*6(DAD =(4( 08*(: 6A;6(4 +@4A:; *6( (:4A'6(+ *6C: *6( :@*4A(:*B)004 )(4A0+
8'%)&0($%*&' O04C9 4((8 =C*(4D @D@C99E '0:*CA: 90= 9(1(9D 08 A:04;C:A' :@*4A(:*DH %6( 4C:;(D 08 :@*4A(:* '0:'(:*4C*A0:D C4( *E)A'C99E, QH- *0 QHX !R :A*4C*(N QHK *0 QHX !R C220:A@2 C:+ 9(DD *6C: QHL !R )60D)604@D C:+ \C:+(4 -5b]?H Y0*6 9C304C*04E C:+ A: DA*@ (G)(4A2(:*D E (* C9H -55Qa \*C239(4 (* C9H -55-a `0>A(9 (* C9H -55P? CD =(99 CD A: DA*@ 03D(41C*A0:D A:D0: (* C9H -55Xa c4A;; -55X?H %6(4( C4( D(1(4C9 =CED 3E =6A'6 (@*40)6A'C*A0: 2CE C+1(4D(9E C7('* '04C9D C:+ '04C9 4((8 DED*(2DH &* *6( ('0DED*(2 9(1(9N :@*4A(:* (:4A'62(:* 2CE (:6C:'( C9;C9 ;40=*6H %6( )409A8(4C*A0: 08 2C'40C9;C( 9(C+D *0 '02B )(*A*A0: 804 D)C'( 3(*=((: C9;C( C:+ '04C9DN =6(4(CD )6E*0)9C:>*0: 39002D A:'4(CD( =C*(4 *@43A+A*E C:+ *6(4(804( +('4(CD( *6( C20@:* 08 9A;6* C1CA9C39( *0 '04C9D 0=D>A (* C9H -55L?H %6( 8@:'*A0:A:; 08 *6( DE23A0DAD 3(*=((: F00GC:*6(99C( C:+ *6(A4 60D* AD *6(4(804( +AD4@)*(+H \02( D*@+A(D 6C1( C9D0 4()04*(+ C '6C:;( A: 'C9'AZB 'C*A0: C:+ ;40=*6 @:+(4 :@*4A(:* (:4A'6(+ '0:+A*A0:DH %6AD 4(D)0:D( AD 1C4AC39( DA:'( 30*6 A:'4(CD(D C:+ +('4(CD(D A: 'C9'AZ'C*A0: 6C1( 3((: 03D(41(+H &* *6( '022@:A*E 9(1(9N \2A*6 C:+ jA:D(E =6(4( : p 5 +@( *0 *6( +(C*6 08 0:( :@33A: +@4A:; *6( (G)(4A2(:*?H &99 *C:>D =(4( C(4C*(+ C:+ 2CA:*CA:(+ @:+(4 A+(:*A'C9 *(2)(4C*@4( D =(4( '9(C:(+ D(1(4C9 *A2(D C =((> A: 04+(4 *0 2A:A2AF( C9;C9 ;40=*6 0: *6( =C99D C:+ :E90: =A4(DH %6( -b =((> A:'@3C*A0: *A2( =CD +A1A+(+ A:*0 80@4 2CA: )(4A0+D, ,3(*%. 8 9,8: ; ? %6AD =CD C )4(B:@*4A(:* (:4A'62(:*

-QX =A*6 C: &99AC:'( VV C@*0C:C9ED(4 80990=A:; *6( 2(*60+ 08 %4(e;@(4 C:+ q( O044( D 9EH &220:A@2 =CD @D(+ CD *6( :A*40;(: D0@4'( DA:'( A* AD *6( 8042 08 :A*40;(: 20D* 4(C+A9E *C>(: @) 3E '04C9D 8402 D(C=C*(4 (4D0: C:+ %4(:'6 -5b.?H O0:'(:*4C*(+ D09@*A0:D 08 /SPO9 C:+ jSK#$P =(4( '0:*A:@0@D9E )@2)(+ A:*0 *6( *C:>D =A*6 C )(4AD*C9*A' )@2)H \(C=C*(4 =CD C9D0 '0:*A:@0@D9E )@2)(+ A: 04+(4 *0 03*CA: Z:C9 / C:+ # '0:'(:*4C*A0:D 08 'CH -Q !R /B/SPh C:+ K_L !R #B#$PL"H %6( D(C=C*(4 r0= 4C*( =CD 'CH QHX 9 6"-H /@*4A(:* '0:'(:*4C*A0:D =(4( 20:A*04(+ (1(4E K_L +CED A: *6( P *C:>DH #60D)604@D '0:'(:*4C*A0:D A: *6( O C:+ / *C:>D =(4( 90= D =(4( -5HX C:+ -bHX !RN 4(D)('*A1(9E D 4(2CA:(+ 90= D =(4( 5 C:+ bHX !RN 4(D)('*A1(9EH V: *6( /# *C:>N :A*40;(: C:+ )60D)604@D '0:'(:*4C*A0:D =(4( D9A;6*9E 90=(4 *6C: A: *6( / C:+ # *C:>DH /A*4C*( C:+ :A*4A*( '0:'(:*4C*A0:D 4(2CA:(+ '0:DAD*(:*9E 90= =CD *0 80990= A:+A1A+@C9 :@33A:D *640@;60@* *6( (G)(4A2(:*C9 )(4A0+D
UC*C C:C9EDAD " !C*(D 08 !#$% N K C:+ J= 804 (C'6 *4(C*2(:* (+N *6( c4((:60@D(Bc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

\0@4'( 08 1C4AC*A0:

\\

+8

R\

M

Y(*=((: D@3k('*D /h# *C:>

#(4A0+D K C:+ L

#(4A0+ P

c p "LHL # -Q"P * h -XH. # -Q"L tt c p "-HP # -Q"P * h .HL # -Q"L tt c p "-H- # -Q"P * h .HX # -Q"L tt

c p LHK # -Q"P * " PLHP # -Q"L tt c p "-HP # -Q"P * h .HL # -Q"L tt c p KHK # -Q"P * " KKHb # -Q"L

@)54$ > %6( =A*6A: D@3k('* 1C4AC*A0:D A: ;40=*6 4C*( =(4( 20+(9(+ A: (C'6 *C:> @DA:; 9A:(C4 4(;4(DDA0:DH !(;4(DDA0: D90)(D =(4( *6(: '02)C4(+ A: (C'6 *C:> 3(*=((: *6( 80@4 +A7(4(:* )(4A0+DN @DA:; C: 04*60;0:C9 '0:*4CD* 6E)0*6(DADH %6( :@99 6E)0*6(DAD =CD (I@C9A*E 08 D90)(DH &9*(4:C*A1( 6E)0*6(D(D =(4(, 6 +('4(CD( A: ;40=*6 4C*( %$J=

=0@9+ 3( @D(+ 804 $K r@G 2(CD@4(2(:*DH O: PN ,N P, A:+A'C*( *6( +A7(4(:* *4(C*2(:*DH Q 0')1>3, W '6C:;( A: 2CGA2C9 ;40DD )60*0DE:*6(*A' 04 ;40=*6 4C*(D +@4A:; )(4A0+ L '02)C4(+ =A*6 )(4A0+ -a ,LK +CA9E ;40DD ,LK 4C*A0

F

K

,LK

#(4A0+ - #(4A0+ L W '6C:;(

#(4A0+ - #(4A0+ L #(4A0+ - #(4A0+L

W '6C:;(

#(4A0+ - #(4A0+ L #(4A0+ - #(4A0+ L

OK O] O5

-. .. LQ

-5 .5 L-

-b L X

LQQ -QX -Q-

LQQ -XQ --X

QHLQ QHPQ QHLX

QHKX QHPL QHLX

"-] ] Q

] L-K

5 LK -P

-H-P -HQ. -HKX

-HQX -HQb -H--

/K /] /5

Pb KQ -]

]K K] -]

XQ LX Q

-QP LQQ -Q-

PQQ PLb LXQ

QHPX QHQ] QHPQ

QHK5 QHQP QH-Q

"L. "XQ "]X

-. -Q b

KQ -L b

-HXQ -HQK -HQ.

-HbQ -HQP -HQ.

#K #] #5

-P L. -b

K] 5K Kb

5L -XX XX

-QK -QX KXQ

LQQ PQK KbQ

QH-] QHPQ QHKQ

QH-Q QHLQ QHQb

"PX "KX "]X

X -. b

] KK -.

-HPQ -H-L -H-K

-H5L KHQ5 QH5Q

/#K /#] /#5

Lb L] XX

.X bQ -QQ

]--. b-

---Kb PQQ

LXQ PQQ PKQ

QH]K QHXL QH]Q

QHKQ QH-] QHKQ

"]L ".b "]K

-b -] KX

KL KK L-

-HQ. -HQ5 -H-Q

-HP-HbK -H.K

XW +@4A:; #PH %6( 9C'> 08 4('01(4E 80990=A:; '(DDC*A0: 08 # (:4A'62(:* 2CE 3( (G)9CA:(+ 3E *6( 8C'* *6C* )60D)604@D C'*D CD C '4ED*C9 )0AD0: ADD -5.P?H V*D (7('* A: *6( D>(9(*0: 2CE *6@D 3( 204( )(4DAD*(:* *6C: *6C* 08 :A*40;(:N =6A'6 C'*D @)0: *6( 204( +E:C2A'N 6A;6B*@4:01(4N 9A1A:; *ADD@( '02)C4*2(:*H S0=(1(4N =6(: )60D)604@D =CD '023A:(+ =A*6 :A*40;(:N C 4C)A+ C:+ D@3D*C:*AC9 4(;CA: A: ;40=*6 =CD 03D(41(+ +@4A:; *6( 4('01(4E )(4A0+H V* D((2D *6(4(804( *6C* )60D)604@D +0(D :0* C'* A: *6( DC2( 2C::(4 =6(: A* AD C90:( CD =6(: A* AD '023A:(+ =A*6 :A*40;(:H O023A:(+ =A*6 :A*40;(:N A*

'C: 3( 4(201(+ 3E C9;C9 ;40=*6 C:+ *6@D 3( )4(1(:*(+ 8402 A:*(48(4A:; =A*6 'C9'AZ'C*A0:H /@*4A(:* (:4A'62(:* 9(+ *0 C: A:'4(CD( A: *6( 4C*( 08 2CGA2C9 ;40DD )60*0DE:*6(DADN =6A'6 =CD @) *0 -XQW 6A;6(4 @:+(4 :@*4A(:*B(:4A'6(+ '0:+A*A0:D
c9E:: C:+ \*(=C4* (4 C:+ $420:+ D C4( +@( *0 /H $@:CAD C:+ D*C7 08 *6( CI@C4A@2 C* *6( $'(C:0;4C)6A' R@D(@2 804 )401A+A:; '04C9DN *0 \H !02CA:(BqA0@+ 804 *('6:A'C9 CDDAD*C:'(N CD =(99 CD *0 UH &99(2C:+N `H O6AD6092 C:+ &H RH \F2C:* 804 6(9)8@9 '022(:*D 0: C: (C49A(4 1(4DA0: 08 *6( )C)(4H %6AD =04> =CD )C4*9E 8@:+(+ 3E *6( #40B ;4C22( /C*A0:C9 !(e'A8D O04C99A(:D A:D0: R` A:D0: R`N OC49D0: YN O40= cq E u (E q! %N \C:+(4 M A:D0: qN SA99 RN dC:; "