全 文 ： ACTA AGRONOMICA SINICA 2008, 34(1): 157−162 http://www.chinacrops.org/zwxb/
ISSN 0496-3490; CODEN TSHPA9 E-mail: xbzw@chinajournal.net.cn

:

 (XJJ2004240)
 :

(1974–), , , ,  !!#$%E-mail: drwanghongwei@sohu.com&
Received (()*): 2007-03-26; Accepted (+,)*): 2007-07-30.
DOI: 10.3724/SP.J.1006.2008.00157
HgCl2  
 1  1  1  2
(1   ; 2   ,  430070)
 :  HgCl2 !#(Vicia faba L.)$% 30 min&, ()*$%+,-./01$%-23456$789
:;<=>? HgCl2-@, )*$%+,-./0@A;B HgCl2CD 10 mg L−1E, FGHIJ#$%
-K:LMN(Pn)6OP:L0QRN(AQY), S<=@A, IJ TAU;E, HgCl2FVWGHXY PSZ:0Q
[0(ΔF/ Fm)6OP:L\Q]^MN(ETR), @A$%-$789:_:`ab(NPQ); cdOe, Y HgCl2fEgF
hi#$% PnXY-jklmnoD HgCl2IJp:L\Q]^q ;
: HgCl2; ;  ;  
Effect of HgCl2 Short Time Treatment on Photosynthesis in Broad
Bean Leaves
WANG Hong-Wei1, SHI Ya-Qi1, HUANG Feng1, and YUAN Lin2
(1 School of Resources and Environmental Engineering, Wuhan University of Technology; 2 Biomedical Materials and Engineering Center,
Wuhan University of Technology, Wuhan 430070, Hubei, China)
Abstract: Mercury pollution is a major environmental problem. Unlike most heavy metals, plants uptake mercury primarily
through leaves rather than the root system. Most of the mercury existed as complexes of Hg2+, which could cause the inhibition of
photosynthesis as well as plant growth in higher plants. Since photosynthesis is very sensitive to heavy metals in higher plants,
green algae and cyanobacteria, the inhibition of photosynthesis by Hg2+ had been examined in CO2 fixation and photosystems
activities. However, in most of the researches, Hg2+ solutions were applied in vitro or not directly on plant leaves, the mechanism
of inhibition on photosynthesis by Hg2+ is still in controversy. In order to discover its effect on leaf photosynthesis, HgCl2 solu-
tions were applied to intact broad bean (Vicia faba L.) leaves. In the experiment, Hg2+ contents in intact broad bean leaves were
determined after 30 min treatment with different concentrations of HgCl2 solution. And leaf gas exchange and chlorophyll fluo-
rescence were measured under a definite light intensity and CO2 concentration. The results showed that the Hg content in leaves
increased with the increase of treated Hg concentrations at the range from 0.5 mg L−1 to 2 000 mg L−1. When the applied HgCl2
concentration was 0.5 mg L−1, the increase of leaf mercury content compared with that of control was less than 20%. If the applied
HgCl2 increased to 500 mg L−1, Hg content in leaves was about 800% of control. Meanwhile, leaf net photosynthetic rate (Pn) was
unaffected at the treated HgCl2 concentrations from 0.5 mg L-1 to 5.0 mg L−1, and gradually decreased with the increase of treated
Hg concentrations from 10 mg L−1 to 2 000 mg L−1. The inhibition of leaf’s Pn was about 36% after 2000 mg L−1 HgCl2 treatment.
The results suggested that broad bean could absorb mercury solution through the leaves and thus lead to the inhibition of leaf
photosynthetic activities.
Further studies showed that both leaf’s apparent quantum yield (AQY) and electron transport rate (ETR) were decreased
significantly, with the increase of HgCl2 concentrations. Moreover, the quantum yield of PS(ΔF/Fm′) were inhibited whereas the
non-photochemical quenching (NPQ) was increased by HgCl2 concentrations from 20 mg L−1 to 2 000 mg L−1, indicating that the
decrease of Pn in broad bean leaves by low concentrations of HgCl2 solutions short time treatment might be mainly due to the
inhibition of photosynthetic electron transport caused by the decrease of light energy transferring to active center and the increase
of the light energy dissipation in PS.
Keywords: Mercuric chloride (HgCl2); Net photosynthetic rate (Pn); Apparent quantum yield (AQY); Chlorophyll fluorescence
158      34

  
[1-2]  !#$, %&
()*+ , ,-*+./012345) , 6
789(:(;<=> , ?@AB(CDE ,
FGHIJK LMN[3]
O PQR(STUVWX, -R( Y=
>Z[4/\]^ _N9`, abR( cd
efghij5[4]klm();<[5-6]nopq
9`rs9`[7-8]
pq9`QR(()tuvwxy./ ,
z{ Hg |pq9`no}~€z‚ƒ
„…, † t‡no CO2 ˆ4[9]; dep#
‰(PS‰)Š‹0Œ [10-11]}no Fd-NADP-Ž
‘’“5[12], ”•–— PS‰“5˜™[13]; 
šnop#›(PS›)“5[14-15]œ, | Hgn
opq9`~ž-Ÿ , ¡¢£ƒQ¤
¥c¦ §¨©(ª. 1 d)|R(«¬, ­©
t‡˜™ƒ®R(-¯p° Pn, ±²³´µ
[7-16]¶·¸[15]¹º[9]»¼½ HgCl2©|¾¿
R(“ ÀÁpq9`«¬ÂąÄ¢
£-©ÅÆ¡ÇÈÉQÊR(ËÌÍÎ- Ï
Ð0ÑQÊ Hg Ò-ÓVR(Ô, —ÕR(
փ ./vw×ØÉÙZ«¬, ‚ÚÛÜ
Hg|pq9`ÝÞ9`~oz„…ßà, Hg|
²³áÀpq9`no‘O-â|R(ËÌã
ä•åÝÞ|pq~æ«¬[16-17]çè, Ä¤
¥ƒé`ê Àë љ¿R(fg9ìí, îÆ
ÛÜ |¾¿R(“ ÀÁpq9`‘9
`ðñ~Oò, óôõ` HgCl2ÏÐö÷“
ÀÁøù»¼½(30 min)©ÅÆ, ¤¥ HgCl2§
¨°úûÀÁpq9`ÀëXüp&ýþ ,
ï€
z{~o
1 
1.1 
úû(Vicia faba L.)âpÓV0(25
/18, 12 h/12 h)[18]R(¡ÌÀÁpuu 300 μmol m−2s−11R(Ÿ ‹, 
[ÀÁ`â HgCl2©, úû
ÀÁÀ9|Ÿ` HgCl2 ÏÐ-
ÀÁ¡°øùö©}| 3, 
£ 50
1.2  
ÀÁ 4!Gp#–,  1 000 W$%
&p, -¯p(  700 μmol m−2s−1 ° 4
Pn
`)*+ ,-Ÿ./ (LI-6400R, LI-COR
01, Lincoln, NE) 4úûÀÁ2pq34[19]
4¼, Õ`56À7(2 cm83 cm), 9p:L;p
q# ?3
500 mL min−1, p( 700 μmol m−2 s−1, CO2=\
400 μmol mo1−1, @AB\ HällgrenÅÆ[20], 4úûÀÁ AQY
1.3 
`CDEq¿ê 9FpG/ Optima 4300
DV ICP-OES ( PerkinElmer Inc. USA) 4úûÀÁ
HuHgCl2 © 30 min $ , ` 10 mmol L−1
Tris-HCl (pH 7.4)IJÀÁ 3KLÀù;M
ê , IJ$NOPQ , R$Ò¤SbTÐ
10 mmol L−1 Tris-HCl (pH 7.4), UŸ¤S$â 4
12 0008g°êŒ 30 min, ¡VÐ, ` 0.22 μmh.
W, & Tyler Olsson [21]ÅÆ 4WÐ Hu
1.4 
& Schreiber¿[22]ÅÆ, )*+ ,-Ÿ
./(LI-6400R, LI-COR01, Lincoln, NE)%X+
üp 4# 4ÀëXüpYZ‹ÀÁ,
4 Fo Fm[ÀÁ- 700 μmol m−2s−1p(°p
q9`©â3!¼, 4 Fm′ Ft 4 Fm′¼, F
ÀÁ 0.8 s¯\p(3 000 μmol m−2s−1)]
°^0+_` PS›apbc4(Fv/Fm)PS›p
u^u(ΔF/Fm′)ødpqCDE34(ETR)
åpbef(NPQ)
ΔF/ Fm′= (Fm′gFt)/ Fm′;
ETR = (Fm′gFt)/ Fm′ × 0.42 × PPFD;
NPQ = (Fm′gFm′)/ Fm′
2 
2.1  HgCl2 !#$ 
%&
¾¿R(hQ.ÀÁ ;< [23]èi
ÕÀÁY HujÒk 1øl, mA HgCl2ÏÐ
=\j , ÀÁYÌ Huno‹jÒ[
HgCl2=\ 0.5 mg L−1¼, ÀÁ Hup|j
Ò
17.22%, qrsÂtZuvw; [ HgCl2=
\jÒ 500 mg L−1¼, ÀÁ Hu 52.5x3.5
μg g−1 Q, Q| 8 yƒ; [=\z{j|
2 000 mg L−1¼, ÀÁ Hu  59.9x1.7 μg g−1
Q
 1 : HgCl2   159




 1  HgCl2  
Fig. 1 Effects of HgCl2 concentrations on leaf mercury content
in broad bean leaves
 3~5  ±
Each value in the figure represent the mean ± SE of 3–5
experiments.

2.2  HgCl2 Pn
HgCl2  30 min,
 700 μmol m−2 s−1 Pn, 
HgCl2   Pn,  !#$
%&&( 2))* HgCl2+, 2 mg
L−1 -, ./012 Pn 345; *6 10 mg
L−1-, Pn7289:, 45;+< 6%)! HgCl2
=>?@, 2 PnABC&, * HgCl2
6 2 000 mg L−1,  D6 36%)
 HgCl2 EF4G,  10
mg L−1 20 mg L−1 HgCl245%&HI CO2
(Ci),J2KLM(GS)NO45 ;  200
mg L−1 HgCl22 CiNO, J45;+<
GS; $B 2 000 mg L−1 HgCl245;+ Ci
GS( 1))PQ Farquhar  Sharkey[24], R
 10~200 mg L−1 HgCl22 PnABSTUV
WXHYZ[;+\KL]; 2 000 mg L−1
HgCl2 EFKL^_, `6ab Pn ;+
cdef)



 2  HgCl2  
Fig. 2 Effects of HgCl2 concentrations on Pn in broad
bean leaves
 6~14  ±,
*** P<0.001
Each value in the table represents the mean ± SE of 6–14 inde-
pendent experiments. ***: Significant at P <0.001.

2.3  HgCl2 AQY
/YghijkYABlZmno
ijfpg, kqrYABZ[s)
thugv(PPFD)w 200 μmol m−2s−11
0 μmol m−2s−1Yxj, y Pn72z{|
}~, €‚ƒ„j6 AQY)
w 2…†‡, HgCl2EF 30 minZˆ45
 AQY)‰Š 20 mg L−1 HgCl2 EF,
  AQY;+< 12%‹Œ)* HgCl2%&
1 200 mg L−1-,   AQY|f;+, E
F628 79%‹Œ; * HgCl2 =>%&
1 2 000 mg L−1-, EFD628 47%‹Œ)Y
Žnghij‘’“h”•–—˜

 1  HgCl2  
Table 1 Effects of HgCl2 concentrations on photosynthetic related parameters in broad bean leaves
 CO2 Ci (μL L−1)

 GS (mmol H2O m−2 s−1)


Concentration
 Control  Treatment  Control  Treatment
10 mg L−1 257.75±0.63 263.33±0.25* 227.22±0.42 224.51±0.61
20 mg L−1 268.79±0.26 277.79±0.21* 322.21±1.71 325.81±1.62
200 mg L−1 279.13±6.38 276.63±6.57 255.82±21.13 230.21±28.01*
2000 mg L−1 271.45±5.49 260.02±5.57* 275.51±22.51 163.22±15.31**
 4  ±, *P<0.05, **P<0.01
Each value in the table represents the mean ± SE of four independent experiments. *Significant at P<0.05, **Significant at P<0.01.

160     ! 34

Y™šn›[[25])dœ,  HgCl2 EFab
 AQY 45;+,ž“h”•Y™šn
›[;)

 2  HgCl2  (AQY)
Table 2 Effects of HgCl2 concentrations on apparent photo-
synthetic quantum yield in broad bean leaves

Concentration
 Control
(mol CO2 mol−1
photons)
 Treatments
(mol CO2 mol−1
photons)
20 mg L−1 0.047#0.005 0.041#0.004**
200 mg L−1 0.046#0.006 0.036#0.003**
2000 mg L−1 0.046#0.005 0.024#0.002***
 4   ±, ** P<0.01,
*** P<0.001
Each value in the table represents the mean±SE of four inde-
pendent experiments.
**: Significant at P<0.01; ***: Significant at P <0.001.

2.4  HgCl2   

Ÿ ¡¢[£k¤¥¦qrY§F
–¨p#©dh2ªX«NOqr›‘
¬C­® , 6¤¥¯°±(PS±)–²Y“h”
•³´<…}µ)¶ 3€G, ‰ 20 200 mg L−1
HgCl2  EF ,  PS±on£ij
(Fv/Fm):28;º , * 2 000 mg L−1 HgCl2  EF , 
Fv/Fm6 0.714, :2845»;< 5 %‹Œ, 
HgCl2 Zˆtf R¯°±¼BZm
nijo½[)Fv/Fm¾n¯°±o
n£ij , Jt¿À8ÁÂ , qrZ
Ã3i»¼BZSTÄÅzÆkΔF/Fm′ (PS±
ghÇg), ȏ<tÉYABÁ
Â, PS±¼BZZ[)‰ 20 mg L−1Ê200 mg
L−1  2 000 mg L−1 HgCl2  EF , 
ΔF/Fm′– ETRËÌ:28;+- NPQ7289:ËÌ%&< 5%Ê8%
31%‹Œ) HgCl2ab  Pn;+ͅ
Zk,È<Y“h”•Š )NPQ
k PS±ÎƒÏ ÐÑZZB,Y“h”•
ÒÓÔÕÖ×Ë, HgCl2  EF NPQ
%&ž HgCl2 EF؁ÙÚZguŠ\n£
ÛÜÕÝÞ<)
3 
fßUž , qrwàálÐÑâ\ã3] ,
Klâkqr‘’âSTUä)åSTk,
àálægç@èéš3êëZ7âÓ`ìµn
Yr, wNOâíî; –wàálÐÑâï
×ËðñtP×, òó×íî[26])ôõ
ö÷øù¤¥ý, , úûlâ3 70.4%~90.7%UV
K, þ3 9.3%~29.6%UVàá[27])qrSTuŠ

‚ÐÑKlâ[23])fßþwqrÓ
¾Í ʖ-»ÄGâ  )
Ú @qr[28-29]ʟ[30-31]X[15,32]
YAB2\ã, Zˆ–-Oz

 3  HgCl2   
Table 3 Effects of HgCl2 concentrations on chlorophyll fluorescence parameters in broad bean leaves

Concentration
PSII
 
Fv/ Fm
PSII 
 
Δ F/ Fm


NPQ

ETR
(μmol m−2 s−1)
20 mg L−1  CK 0.775±0.005 0.495±0.006 0.282±0.003 145.628±1.604

 TR 0.750±0.010 0.446±0.005 0.296±0.005 131.036±1.444

/ TR/CK (%) 96.77 90.10** 104.96 89.98**

200 mg L−1  CK 0.764±0.010 0.486±0.006 0.290±0.007 142.982±1.799

 TR 0.731±0.010 0.404±0.004 0.314±0.005 118.678±1.251

/ TR/CK (%) 95.68 83.13*** 108.28* 83.00***

2000 mg L−1  CK 0.748±0.009 0.459±0.011 0.289±0.004 134.848±3.113

 TR 0.714±0.006 0.331±0.010 0.381±0.008 97.314±2.984

/ TR/CK (%) 95.45* 72.11*** 131.83*** 72.17***
!#$ 4%&()*+,-±./01* 2 P<0.05, ** 2 P<0.01, *** 2 P<0.0011
Each value in the table represents the mean ± SE of four independent experiments. *: Significant at P<0.05; **:Significant at P<0.01; ***:
Significant at P <0.001.
 1 : HgCl2   161


ª¾n)d¤¥â2YABN
O6³´Ú¤¥fâZˆqr
YAB, Jt²êF
!tË#)$%&6â
Z(ë)*9+AB[33], t“h”•,ë
 [34]Ú-./E01“h”•, 2¤¥6¦
3 HgCl2 YABê³´<456)
’×âæg7 Hg2+|8qr9:XH, 
, ç+ HgCl2EF,  Pn;+ST
k,<Y“h”•Š ;\<
1KL]; @(=Ìk 2 000 mg L−1)HgCl2E
F, KL^_Y“h”•Š ·<1
, >` Pn;+cd?@AKLBCd
 )J, 2 000 mg L−1HgCl2EF2 AQY
ETR ABç, Pn;+DD6 36%, …
Eœ-\KLd kabYAB;STc
d)!âFh³@, KL^_Y“h
”•< %&, =ÌkG2 HgCl2 E
FÙ)‰H-EF, WXHYê«
:IJXHÙKL<1 HgCl2 NO, MGIJX
HtWXH Pn ;NOP)Q¿RÚF‘Ÿ‘¤¥ HgCl2 2“h
”•NO, 2SRt›‘¹ºTG<WXH
Yê«“h”•¯°…Zk HgCl2 YA
BSTUVAB./)*W3Bå
¤¥
!XT|fgYA¦3)
-, !Z[qr2âÐÑ#©
lâ3fB¯ , t92ç+ , qr
2 Hg ÐÑj92ç@)å7 Stratton [35]
* HgCl2 6 2 mg L−1-, \]^_
(Anabaena inaequalis)2 HgCl2ÐÑZˆ¸1 96%
f`)2SRH-IEF, âFha
9:ÐÑ:bDt 0.4%~13.5%eI)t92
ç@#$ HgCl2, qr2 HgÐÑj
92ç+; t92ç+, qr2 Hg
ÐÑj92ç@)dœ, ZcdeØKl
t+Kâ, qr?…gÐÑf
gâ…Zµ)▲nYrÍaqr;hMm
n6ijÓÔ, ÙÚkl, ‘’fg Hg!mm
n62§r‘Ù3no*â)€, pqdâ
tArlÐÑÊfg–2YABNO
rs)
4 
 HgCl2 30 minEF,  
 Pn AQY t3 ;+, ;+Du
v,#$ , -,   ΔF/Fm
ETR 45;+, NPQ %&)HgCl2ab  Pn
;+cdtÍ Rk<Y“h”•Š
, ΃ϠòABlw”•Zxy PS±Ö
ÕZ%&)
References
[1] Schroeder W H, Munthe J. Atmospheric mercuryan over-
view. Atmos Environ, 1998, 32: 809−822
[2] Boening D W. Ecological effects, transport and fate of mer-
cury: A general review. Chemosphere, 2000, 40: 1335−1351
[3] Suszeynsky E M, Shann J R. Phytotoxicity and accumulation
of mercury in tobacco subjected to different exposure routes.
Environ Toxicol Chem, 1995, 14: 61−67
[4] Ma C-C(). Hg harm on cell membrane of rape leaf and
cell endogenous protection effect. Chin J Appl Ecol (
), 1998, 9(3): 323−326 (In Chinese with English ab-
stract)
[5] Cho U-H, Park J-O. Mercury induced oxidative stress in to-
mato seedlings. Plant Sci, 2000, 156: 1−9
[6] Patra M, Sharma A. Mercury toxicity in plants. Bot Rev, 2000,
66: 379−422
[7] Godbold D L, Huttermann A. The uptake and toxicity of
mercury and lead to spruce (Picea abies Karst.) seedlings.
Water Air Soil Poll, 1986, 31(1/2): 509−515
[8] Krupa Z, Baszynski T. Some aspects of heavy metals toxicity
towards photosynthetic apparatus: direct and indirect effects
on light and dark reactions. Acta Physiol Plant, 1995, 17(2):
177−190
[9] De Filippis L F, Ziegler H. Effect of sublethal concentrations
of zinc, cadmium and mercury on the photosynthetic carbon
reduction cycle of Euglena. J Plant Physiol, 1993, 142:
167−172
[10] Golbeck J H, Lein S, San Pietro A. Isolation and characteri-
zation of a sub-chloroplast particle enriched in iron sulfur
protein and P700. Arch Biochem Biophys, 1977, 178: 140−150
[11] Kojima Y, Hiyama T, Sakurai H. Effect of mercurials on iron
sulfur centers of PSof Anacystis nidulans. In: Biggins J ed.
Progress in Photosynthesis Research. Dordrecht: Martinus
Nijhoff, 1987. pp 57−60
[12] Honeycutt R C, Krogmann D W. Inhibition of chloroplast re-
actions with phenylmercuric acetate. Plant Physiol, 1972, 49:
376−380
[13] De Filippis L F, Hampp R, Ziegler H. The effects of sublethal
concentrations of zinc, cadmium and mercury on Euglena: .
Respiration, photosynthesis and photochemical activities.
Arch Microbiol, 1981, 128: 407−411
[14] Murthy S D S, Sabat S C, Mohanty P. Mercury-induced inhi-
bition of photosystem activity and changes in the emis-
sion of fluorescence from phycobilisomes in intact cells of the
162      34

cyanobacterium, Spirulina platensis. Plant Cell Physiol, 1989,
30: 1153−1157
[15] Singh D P, Khare P, Bisen P S. Effects of Ni2+, Hg2+ and Cu2+
on growth, oxygen evolution and photosynthetic electron
transport in Cylindrospermum IU 942. J Plant Physiol, 1989,
134: 406−412
[16] Godbold D L. Mercury-induced root damage in spruce seed-
lings. Water Air Soil Poll, 1991, 56: 823−831
[17] Godbold D L, Huettermann A. Effect of zinc, cadmium and
Hg on root elongation Picea (Karst) seedlings and the sig-
nificance of these metals to forest die-back. Environ Poll (ser
A), 1985, 38(4): 375−381
[18] Yuan L, Xu D Q. Stimulation effect of gibberellic acid
short-term treatment on leaf photosynthesis related to the in-
crease in Rubisco content in broad bean and soybean. Photo-
synth Res, 2001, 68(1): 39−47
[19] Wang H W, Su J H, Shen Y G. Differences in response of
photosynthesis to bisulfite between two wheat genotypes. J
Plant Physiol Mol Biol, 2003, 29: 27−32
[20] Long S P, Hällgren J E. Measurement of CO2 assimilation by
plants in the field and the laboratory. In: Techniques in Bio-
productivity and Photosynthesis. Oxford: Pergamon Press,
1985. pp 62−94
[21] Tyler G, Olsson T. The importance of atmospheric deposition,
charge and atomic mass to the dynamics of minor and rare
elements in developing, ageing, and wilted leaves of beech
(Fagus sylvatica L.). Chemosphere, 2006, 65: 250−260
[22] Schreiber U, Bilger W, Hormann H, Neubauer C. Chlorophyll
fluorescence as a diagnostic tool: basics and some aspects of
practical relevance. In: Raghavendra A S ed. Photosynthesis:
A Comprehensive Treatise. Cambridge: Cambridge University
Press, 1998. pp 320−336
[23] Lindberg S E, Kim K-H, Meyers T P, Owens J G. Micromete-
orological gradient approach for quantifying air/surface ex-
change of mercury vapor. Environ Sci Tech, 1995, 29:
126−135
[24] Farquhar G D, Sharkey T D. Stomatal conductance and pho-
tosynthesis. Annu Rev Plant Physiol, 1982, 33: 317−345
[25] Foyer C H, Harbinson J. The photosynthetic electron transport
system: efficiency and control. In: Foyer C H, Quick W P eds.
A Molecular Approach to Primary Metabolism in Higher
Plants. London: Taylor & Francis, 1997. pp 3−39
[26] Fu X-W( ), Feng X-B(), Wang S-F(), Li
Z-G (), Li P(). Advances of research on mercury
in plants. Bull Miner Petrol Geochem( ! #
), 2005, 24(3): 232−238 (in Chinese with English abstract)
[27] Liu D-S($%&), Qing C-L(()). Contribution to vegeta-
ble mercury from atmosphere and soil. Chin J Appl Ecol(
), 2002, 13(3): 315−318 (in Chinese with English ab-
stract)
[28] Clijsters H, van Assche F. Inhibition of photosynthesis by
heavy metals. Photosynth Res, 1985, 7(1): 31−40
[29] Sheoran I S, Singh R. Effect of heavy metals on photosynthe-
sis in higher plants. In: Abrol Y P, Mohanty P, Govindjee eds.
Photosynthesis: Photoreactions to Plant Productivity.
Dordrecht: Kluwer Academic Publishers, 1993. pp 451−468
[30] Shioi Y, Tamai H, Sasa T. Inhibition of photosystem in
the green alga Ankistrodesmus falcatus by copper. Physiol
Plant, 1978, 44: 434−438
[31] Samson G, Morisette J C, Popovic R. Copper quenching of the
variable fluorescence in Dunaliella tertiolecta. New evidence
for a copper inhibition effect on PSII photochemistry. Photo-
chem Photobiol, 1988, 48: 329−332
[32] Singh D P, Singh S P. Action of heavy metals on Hill activity
and O2 evolution in Anacystis nidulans. Plant Physiol, 1987,
83: 12−14
[33] Passow H, Rothstein A, Clarkson T W. The general pharma-
cology of heavy metals. Pharmacol Rev, 1961, 13:185−224
[34] Katoh S, Takamiya A. Nature of copper protein binding in
spinach plastocyanin. J Biochem, 1964, 55: 378−387
[35] Stratton G W, Huber A L, Corke C T. Effect of mercuric ion
on the growth, photosynthesis, and nitrogenase activity of
Anabaena inaequalis. Appl Environ Microbiol, 1979, 38(3):
537−543