563
Photosynthesis in Nature: A New Look
by higher plants and algae in photosynthesis is originated from H 2 O and not from CO 2
(Ruben et al. 1941; Vinogradov and Teı ˇs 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 during 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 formation 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 proposed to react with CO 2 in photosynthesis, whereas H 2 O 2 is used as an intermediate 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,
Photosynthesis in Nature: A New Look
by higher plants and algae in photosynthesis is originated from H 2 O and not from CO 2
(Ruben et al. 1941; Vinogradov and Teı ˇs 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 during 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 formation 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 proposed to react with CO 2 in photosynthesis, whereas H 2 O 2 is used as an intermediate 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,