English (pdf) - SciELO

5*146 %1//70+%#6+10

5VWF[ QH VJG GHHGEV QH #NKGVVG QP VJG CEVKXKV[ QH URKPCEJ

5RKPCEGC QNGTCEGC . EJNQTQRNCUVU 2GFTQ#NOGKFC,QºQ2%QWVKPJQ,QºQ4QEJCCPF,QUÃ)QOGU.CTCPLQ %GPVTQFG'UVWFQU6GEPQNÎIKEQUFQ#ODKGPVGGFC8KFC7PKXGTUKFCFG6T¶UQU/QPVGUG#NVQ&QWTQ8KNC 4GCN 2QTVWICN %QTTGURQPFKPI CWVJQT LNCTCPLQ"WVCFRV 4GEGKXGF,CPWCT[4GVWTPGFHQTTGXKUKQP#RTKN#EEGRVGF,WPG

+PVGPUKHKECVKQP QH RTQFWEVKQP KP OQFGTP CITKEWNVWTG KU HTGSWGPVN[ QDVCKPGF VJCPMU VQ VJG KPETGCUG QH RGUVKEKFG CRRNKECVKQPU CPF EJGOKECN HGTVKNK\GTU *QYGXGT VJG WUG QH VJGUG RTQFWEVU UQOGVKOGU ECWUGU C TGFWEVKQP QH ETQR RTQFWEVKXKV[9GCKOGFVQGXCNWCVGVJGKORCEVQHVJGHWPIKEKFG#NKGVVGQPKPXKVTQCEVKXKV[QHEJNQTQRNCUVUQHURKPCEJ

5RKPCEGCQNGTCEGC.6TCPUOGODTCPGRQVGPVKCNHQTOCVKQP25++QZ[IGPGXQNWVKQP CPF25++RJQVQEJGOKUVT[YGTG CPCN[\GF 1WT TGUWNVU UJQYGF VJCV VJG TGEQOOGPFGF EQPEGPVTCVKQP  I .  QH #NKGVVG FKF PQV KORCKT VJG RJQVQU[PVJGVKECRRCTCVWU*QYGXGTCVJKIJGTEQPEGPVTCVKQPU I. TGFWEVKQPUKPVJ[NCMQKFOGODTCPGRQVGPVKCN CPFQZ[IGPGXQNWVKQPYGTGTGEQTFGFDWVYKVJPQGXKFGPEGQHRJQVQKPJKDKVKQP -G[YQTFUDKQGPGTIGVKEUGEQVQZKEQNQI[RGUVKEKFGURJQVQU[PVJGUKUZGPQDKQVKEU

# KPVGPUKHKECÁºQ FC RTQFWÁºQ PC CITKEWNVWTC OQFGTPC à QDVKFC OWKVCU XG\GU ITCÁCU CQ KPETGOGPVQ FG CRNKECÁÐGU FG RGUVKEKFCU G HGTVKNK\CPVGU OKPGTCKU 0Q GPVCPVQ Q WUQ FGUUGU RTQFWVQU ECWUC RQT XG\GU C TGFWÁºQ FC RTQFWÁºQ 1 QDLGVKXQFGUVGGUVWFQHQKCPCNKUCTKPXKVTQQGHGKVQFQHWPIKEKFC#NKGVVGUQDTGCCVKXKFCFGFGENQTQRNCUVQUFGGURKPCHTG

5RKPCEGCQNGTCEGC.1UTGUWNVCFQUOQUVTCTCOSWGCEQPEGPVTCÁºQVKRKECOGPVGTGEQOGPFCFC I.FG#NKGVVGPºQ CHGVQWPGICVKXCOGPVGQCRCTGNJQHQVQUUKPVÃVKEQ0QGPVCPVQCEQPEGPVTCÁÐGUOCKUGNGXCFCU I. TGIKUVTCTCOUG TGFWÁÐGUPQRQVGPEKCNFGOGODTCPCFQUVKNCEÎKFGUGPCRTQFWÁºQFGQZKIÄPKQRQTÃOUGOGXKFÄPEKCFGHQVQKPKDKÁºQ 2CNCXTCUEJCXGDKQGPGTIÃVKECGEQVQZKEQNQIKCHQVQUUÈPVGUGRGUVKEKFCUZGPQDKÎVKEQU

9KVJVJGVTGOGPFQWUFGXGNQROGPVQHCITKEWNVWTGCHVGTVJG GPF QH 9QTNF 9CT ++ VJG GZRTGUUKQP ő/QFGTP #ITKEWNVWTGŒYCUFGXGNQRGFCUQRRQUGFVQVJGEQPEGRV QHő6TCFKVKQPCN#ITKEWNVWTGŒ$GHQTGVJGYCTVJKUYCUVJG QPN[ V[RG QH CITKEWNVWTG RTCEVKUGF 6JKU PGY MKPF QH CITKEWNVWTG DTQWIJV VJG PGGF HQT NCTIGT CTGCU QH JGCX[ OCEJKPGU CPF VJG WUG QH EJGOKECN RTQFWEVU YKVJ VJG CDKNKV[ VQ EQPVTQN VJG FGXGNQROGPV QH UQOG QTICPKUOU EQPUKFGTGF CU KPHGEVCPVU 9QNHH  UQOGVKOGU FKUTGURGEVKPI GPXKTQPOGPVCN TWNGU *QYGXGT UWEJ EJGOKECNU ECP ECWUG UGXGTG GPXKTQPOGPVCN RTQDNGOU UKPEG VJG[ CTG QHVGP WUGF CV EQPEGPVTCVKQPU JKIJGT VJCP VJQUG TGEQOOGPFGF QWV QH UGCUQP QT KP EWNVWTGU HQT YJKEJVJG[YGTGPQVQTKIKPCNN[FGXGNQRGFPQVVQOGPVKQP VJG WUG QH HQTDKFFGP RTQFWEVU +P HCEV UWEJ EJGOKECNU

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

#ITQ $C[GT  #NKGVVG KU C DKFKTGEVKQPCN U[UVGOKE HWPIKEKFG $TQYP  YJKEJ CEVU FKTGEVN[ QP HWPIK

$TC\,2NCPV2J[UKQN 

2#./'+&# GV CN

UVQRRKPIVJGITQYVJQHO[EGNKCTGFWEKPIVJGRTQFWEVKQP QH URQTGU CPF TGKPHQTEKPI VJG PCVWTCN FGHGPEGU QH VJG RNCPV #ITQ $C[GT  6JKU RTQFWEV JCU PQ VQZKEQNQIKECN ENCUUKHKECVKQP DWV VJG FQUG QH  I .  CRRNKGF QP HQNKCIG KU TGEQOOGPFGF CICKPUV HWPIK NKMG 2J[VJKWO CPF 2J[VQRJVJQTC VJCV CHHGEV UGXGTCN ETQRU UWEJCUEKVTWUNGVVWEGCPFJQR 1TOGCPF-GING[ +V KU CNUQ WUGF HQT EJGUVPWV VQ NKOKV KPM FKUGCUG KORCEV ECWUGFD[2J[VQRJVJQTCEKPPCOQOK )WGUVGVCN %JGUVPWVEWNVKXCVKQPJCUGZRGTKGPEGFCNCTIGGZRCPUKQPKP 6T¶UQU/QPVGU 2QTVWICNYJKEJTGRTGUGPVUCNOQUV QH2QTVWICNŏ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

%JCPIGUKPVJ[NCMQKFOGODTCPGRQVGPVKCN '< YGTGQDUGTXGFYKVJCHNWQTGUEGPVECVKQPKERTQDGCOKPQ EJNQTQOGVJQZ[CETKFKPG #%/#CEEQTFKPIVQ)Q OGU.CTCPLQGVCN +PVJGEQPVTQNCUUC[ P/QH#%/#YGTGCFFGFCHVGTOKPVQIEJNQTQRJ[NN OO/UQTDKVQNO/VTKEKPG0C1* R*O/ /I%NO/-%NCPFO/OGVJ[NXKQNQIGP#HVGT OKP VJG EJNQTQRNCUVU YGTG GZEKVGF YKVJ TGF TCFKCVKQP

POO9EORTQXKFGFHTQODGNQYHQTOKP+P VJG CUUC[U YKVJ #NKGVVG VJG QPN[ FKHHGTGPEG YCU VJCV DGHQTGVJGCFFKVKQPQH#%/##NKGVVGYCUCFFGFKPXKVTQ IKXKPI C HKPCN EQPEGPVTCVKQP DGVYGGP  CPF  I .  1Z[IGP GXQNWVKQP YCU UKOWNVCPGQWUN[ OGCUWTGF RQNCTQITCRJKECNN[ WUKPI C %NCTMV[RG QZ[IGP GNGEVTQFG

%CTRGPVKGTGVCN %JNQTQRJ[NN C HNWQTGUEGPEG YCU OQPKVQTGF YKVJ CP (/5 HNWQTQOGVGT *CPUCVGEJ 0QTHQNM 7- 6JG TGCEVKQPOGFKWOYCUVJGUCOGCUVJCVWUGFKPVJGUVWFKGU YKVJ #%/# /GCUWTGOGPVU YGTG OCFG QH DCUCN HNWQTGUEGPEG (   OCZKOCN HNWQTGUEGPEG ( O WPFGT $TC\,2NCPV2J[UKQN 

UCVWTCVKPINKIJVRWNUG zOQNO UUVGCF[UVCVG HNWQTGUEGPEG (UCVCEVKPKETCFKCVKQP POQNOUCPF OCZKOCN HNWQTGUEGPEG YKVJ TGCEVKQP EGPVTGU RCTVKCNN[ ENQUGF (Oŏ (KIWTG  (TQO VJGUG RCTCOGVGTU ( X( O

XCTKCDNGVQOCZKOCNHNWQTGUEGPEGTCVKQYJKEJTGRTGUGPVU VJG OCZKOCN RJQVQEJGOKECN GHHKEKGPE[ QH 25++ ) 25++

SWCPVWO [KGNF QH 25++ GNGEVTQP VTCPURQTV S R

RJQVQEJGOKECN SWGPEJKPI EQGHHKEKGPV CPF S 02 PQP RJQVQEJGOKECN SWGPEJKPI EQGHHKEKGPV YGTG ECNEWNCVGF CEEQTFKPIVQ.CKUMCPF1LC #NNTGUWNVURTGUGPVGF JGTG YGTG HTQO C UKPING CUUC[ DWV UKOKNCT TGUWNVU YGTG QDVCKPGFKPVJTGGKPFGRGPFGPVGZRGTKOGPVU#PCN[UGUQH FCVCYGTGECTTKGFQWVWUKPIVJG/KETQUQHV'ZEGNRTQITCO KP QTFGT VQ QDVCKP VJG UGEQPFFGITGG RQN[PQOKCN TGITGUUKQPOQFGNU 120

Fluorescence (mV)



A

100

C

Fm

80 Fv

60

B

Control 20 mM

Fv'

40 20 0 0

20

40

60 Time (s)

80

100

(KIWTG  6[RKECN VTCEG QH EJNQTQRJ[NN HNWQTGUEGPEG QDVCKPGFYKVJHTGGZGPQDKQVKEURKPCEJEJNQTQRNCUVGZVTCEV CPFCHVGTCRRNKECVKQPQHO/#NKGVVG$CUCN (OCZKOCN

(OCPFXCTKCDNG (XHNWQTGUEGPEGYGTGFGVGTOKPGFKPFCTM CFCRVGF UCORNGU QRGP 25++ TGCEVKQP EGPVTGU ( O YCU QDVCKPGFCHVGTCPCEVKPKENKIJVRWNUG #+PNKIJVCFCRVGF UCORNGUUVGCF[UVCVG (UOCZKOCN (OŏCPFXCTKCDNG (Xŏ HNWQTGUEGPEG YGTG OGCUWTGF WPFGT FKO NKIJV CHVGT $ EQPFKVKQPUYKVJRCTVKCNN[QRGP25++TGCEVKQPEGPVTGUCPF CHVGTCPGYCEVKPKENKIJVRWNUG % +V UJQWNF DG PQVGF VJCV VJG TGEQOOGPFGF #NKGVVG EQPEGPVTCVKQP VQ DG CRRNKGF QP HQNKCIG KU  I .   YJGTGCU KP VJKU UVWF[ VJG HWPIKEKFG YCU CRRNKGF CV EQPEGPVTCVKQPU TCPIKPI HTQO  VQ  I .  FKTGEVN[ VQ C VJ[NCMQKF UWURGPUKQP 5KPEG QPN[ C HTCEVKQP QH #NKGVVG CRRNKGFQPHQNKCIGTGCEJGUVJGRJQVQU[PVJGVKECRRCTCVWU CPGZCEGTDCVKQPQHRQVGPVKCNGHHGEVUQHVJGHWPIKEKFGQP RJQVQU[PVJGUKU HQT C IKXGP EQPEGPVTCVKQP KU VQ DG GZRGEVGFKPVJKUUVWF[KPEQORCTKUQPYKVJHQNKCT#NKGVVG CRRNKECVKQPU+PCP[ECUGQWTTGUWNVUUJQYGFVJCV#NKGVVG



'(('%6 1( #.+'66' 10 6*' #%6+8+6; 1( %*.1412.#56

A

110 Fluorescence (% control)

100 90 80 70 60

t0.51,t0.5d (s)

0

2.5

5

7.5 10 12.5 15 17.5 20 22.5

16 14 12 10 8 6 4 2 0

t0.5d

B

t0.5l

0

Oxygen evolution (% control)

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

2.5

5

7.5

10 12.5 15 17.5 20 22.5

C

110 100 90 80 70 60 50 0

2.5

5

7.5 10 12.5 15 17.5 20 22.5 Aliette (g L-1)

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

FCVCPQVUJQYPYJKEJKUUKOKNCTVQVJCVQDVCKPGFHQTVJG TGEQOOGPFGF EQPEGPVTCVKQP YKVJ JKIJGT #NKGVVG EQPEGPVTCVKQPU C NQPIGT VKOG YCU PGGFGF VQ TGCEJ VJCV RQVGPVKCN 6JCV EQWNF DG FWG VQ C FGITCFCVKQP QH VJG

$TC\,2NCPV2J[UKQN 



2#./'+&# GV CN

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ŏUKPEGPQUKIPKHKECPVXCTKCVKQPYCU QDUGTXGFKP(X (KIWTG#EEQTFKPIVQ-TCWUGCPF9GKU

VJKUSWGPEJKPIEQWNFDGFWGVQVJGFKOKPWVKQPQH VJG KPVTCVJ[NCMQKF CEKFKHKECVKQP ECWUGF D[ GNGEVTQP VTCPURQTVYJKEJYCUCNUQFGOQPUVTCVGFD[VJGTGFWEVKQP QHVJGOGODTCPGRQVGPVKCN (KIWTG# (TQO VJG CDQXG YG EQPENWFG VJCV #NKGVVG CV VJG TGEQOOGPFGF EQPEGPVTCVKQP FQGU PQV KORCKT VJG HWPEVKQPKPI QH VJG RJQVQU[PVJGVKE OCEJKPGT[ DWV CV EQPEGPVTCVKQPU CDQXG VJCV TGEQOOGPFGF KV ECP PGICVKXGN[ KORCEV VJG DKQGPGTIGVKEU QH URKPCEJ EJNQTQRNCUVU

4'('4'0%'5 (KIWTG'HHGEVQH #NKGVVGCVUGXGTCNEQPEGPVTCVKQPUQP EJNQTQRJ[NNCHNWQTGUEGPEGRCTCOGVGTUGXCNWCVGFKPURKPCEJ EJNQTQRNCUVU XCTKCDNGVQOCZKOCN FCTMCFCRVGF HNWQTGUEGPEG TCVKQ ( X( O # SWCPVWO [KGNF QH 25++ GNGEVTQP VTCPURQTV )25++ $ RJQVQEJGOKECN S R % CPF PQPRJQVQEJGOKECNS02 &SWGPEJKPIEQGHHKEKGPVU5GG NGIGPFVQ(KIWTGHQTQVJGTFGVCKNU

$TC\,2NCPV2J[UKQN 

#ITQ$C[GT 2TQRTKGFCFGU#NKGVVG(NCUJ #EEGUU JVVRYYYCITQDC[GTEQORV $TQYP 5  +PUGPUKVKXKV[ VQ VJG HWPIKEKFG HQUGV[N CNWOKPKWOKP%CNKHQTPKCKUQNCVGUQHVJGNGVVWEGFQYP[ OKNFGYRCVJQIGP$TGOKCNCEVWECG2NCPV&KU  %CTRGPVKGT4.CTWG$.GDNCPE4/ 2JQVQCEQWUVKE URGEVTQUEQR[QH#PCE[UVKUPKFWNCPU+++&GVGEVKQPQH

'(('%6 1( #.+'66' 10 6*' #%6+8+6; 1( %*.1412.#56

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