Environmental Engineering Reference
In-Depth Information
by higher plants and algae in photosynthesis is originated from H
2
O and not from CO
2
(Ruben et al.
1941
; Vinogradov and Teˇs
1941
).
Calvin and his colleagues (A. Benson and J. Bassham) during the period of
1947-1956 depicted the mechanism that carbon travels through a plant dur-
ing photosynthesis, starting from its absorption as atmospheric carbon dioxide to
its conversion into carbohydrates and other organic compounds (Bassham et al.
1956
; Calvin
1956
). They showed that sunlight can act on chlorophyll in a plant to
fuel the production of organic compounds, rather than on CO
2
as was previously
believed. Calvin was then awarded the Nobel Prize in Chemistry in 1961 for his
Calvin cycle (sometimes termed as Calvin-Benson-Bassham Cycle).
Since then, a lot of studies have been conducted on photosynthesis regarding
release of electrons from chlorophylls, characterization of the primary reaction
centers of photosystems (PSI and PSII), occurrence and formation of chlorophyll
dimer in PSI and PSII, functions and roles of PSI and PSII, endogenous forma-
tion of hydrogen peroxide (H
2
O
2
) in photosynthetic cells, release of O
2
, and so
on (Hill
1937
,
1939
; van Niel
1931
; Bach
1894
; Arnon
1949
,
1959
,
1961
,
1971
;
Mehler
1951
; Asada et al.
1974
; Chance et al.
1979
; Halliwell
1981
; Holland
et al.
1982
; Boekema et al.
1987
,
2001
Shipman et al.
1976
; Hynninen and
Lötjönen
1993
; Krauss et al.
1993
; Krauß et al.
1996
; Shubin et al.
1993
; Golbeck
1994
; Kruip et al.
1994
; Boussaad et al.
1997
; Brettel
1997
; Wilhelm et al.
1996
,
1997
,
1999
; Klukas et al.
1999
; Halliwell and Gutteridge
1999
; López-Huertas
et al.
1999
; Stewart et al.
2000
; Jordan et al.
2001
; Baker and Graham
2002
;
Ben-Shem et al.
2003
; Catalan et al.
2004
; Dashdorj et al.
2004
; Germano et al.
2004
; Diner and Rappaport
2002
; del Río et al.
2006
; Li et al.
2006
; Krasnovsky
2007
; Krieger-Liszkay et al.
2008
; Rappaport and Diner
2008
; Amunts et al.
2010
;
Müller et al.
2010
; Nilsson Lill
2011
; Umena et al.
2011
).
Moreover, release of O
2
during photosynthesis still remains under debate
because it is considered to be originated either from H
2
O (Dashdorj et al.
2004
;
Germano et al.
2004
; Rappaport and Diner
2008
; Müller et al.
2010
; Takahashi
et al.
1987
; Periasamy et al.
1978
) or from H
2
O
2
(Komissarov
1994
,
2003
; Bach
1893
; Velthuys and Kok
1978
; Asada and Badger
1984
; Asada and Takahashi
1987
; Mano et al.
1987
; Renger
1987
; Anan'ev and Klimov
1988
; Bader and
Schmid
1988
,
1989
; Schroeder
1989
; Schröder and Åkerlund
1990
; Miyake and
Asada
1992
; Bader
1994
,
1995
; Yin et al.
2006
; Mostofa et al.
2009
; Kuznetsov
et al.
2010
; Bernardini et al.
2011
; Mostofa et al.
2009
). The scientist Bach has
been the first to show that plants actively accumulate H
2
O
2
upon illumination
(Bach
1894
). The major generation sites of reactive oxygen species (ROS) are
the PSI and PSII photosystems in chloroplast thylakoids in higher plants. Here,
photoreduction of O
2
to H
2
O
2
in PSI has firstly been discovered by Mehler
(Mehler
1951
). Subsequently, the primary reduced product has been identified
as the superoxide anion (O
2
•
−
), the disproportionation of which can produce
H
2
O
2
and O
2
(Asada et al.
1974
). Recently, H
2
O
2
instead of H
2
O has been pro-
posed to react with CO
2
in photosynthesis, whereas H
2
O
2
is used as an inter-
mediate to release O
2
(Komissarov
1994
,
2003
; Velthuys and Kok
1978
; Mano
et al.
1987
; Renger
1987
; Anan'ev and Klimov
1988
; Bader and Schmid
1988
,