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high-temperature episodes when leaf temperature can exceed the air temperature
by as much as 15 °C (Singsaas and Sharkey 1998 ; Hanson et al. 1999 ; Singsaas et
al. 1999 ). Rubisco can produce hydrogen peroxide as a result of oxygenase side
reactions, which can increase substantially with temperature (Sharkey 2005 ).
Moreover, an increase in temperature can induce sinks of electron transport
different from CO 2 assimilation, and photorespiration is increased at 30-35 °C
(D'Ambrosio et al. 2006 ). The O 2 -independent electron transport can account for up
to 20 % of the total PSII electron transport in wild watermelon leaves (Miyake and
Yokota 2000 , 2001 ). The electron flux in PSII that exceeds the flux required for the
cycles of photosynthetic carbon reduction and photorespiratory carbon oxidation,
can induce photoreduction of O 2 in the water-water cycle (Miyake and Yokota 2000 ,
2001 ). It has been shown that the greater partitioning of reductive power to non-
assimilative processes consuming O 2 (photorespiration, Mehler reaction and chlo-
rorespiration) with respect to CO 2 assimilation allows keeping the PSII efficiency
factor unmodified at temperatures as high as 35 °C (D'Ambrosio et al. 2006 ).
The unsaturation of fatty acids can protect PSII from the inhibition of the activ-
ity that is caused by strong light at low temperatures (Wada et al. 1990 ; Murata
et al. 1992 ), and can accelerate the repair of photodamaged PSII (Gombos
et al. 1994 ; Wada et al. 1994 ; Moon et al. 1995 ). After photodamage to PSII in
Synechocystis at low temperatures (0-10 °C), activity recovery can reach up to
50 % of the original level in the darkness at moderate temperatures, without the de
novo synthesis of D1 protein (Nishiyama et al. 2008 ).
High-temperature stress can disrupt the cellular metabolic homeostasis and promote
the production of reactive oxygen species (H 2 O 2 , 1 O 2 , O 2
) (Mittler 2002 ).
Oxidative stress occurs in any plant cell when there is an imbalance between produc-
tion of ROS and antioxidant defense (Apel and Hirt 2004 ; Mittler 2002 ; Scandalios
2002 ). The consequence is a decrease of the net photosynthetic efficiency that affects
various plant activities (Ogweno et al. 2008 ; Apel and Hirt 2004 ; García-Ferris and
Moreno 1994 ; Alscher et al. 1997 ; Anderson 2002 ; Irihimovitch and Shapira 2000 ;
Pfannschmidt 2003 ). Calvin-cycle enzymes within chloroplasts are particularly sensi-
tive to high levels of H 2 O 2 , which decreases CO 2 fixation and foliar biomass (Willekens
et al. 1997 ; Zhou et al. 2004 , 2006 ). The mechanism behind the decline of plant photo-
synthesis by high-temperature stress, driven by high irradiance or drought or heat stress,
is similar to that of high irradiance as mentioned earlier.
, and HO
5.4 Effects of Water Stress (Drought)
and of Precipitation/Rainfall
Water stress or drought stress can significantly affect plant photosynthesis and
decrease their growth, development and productivity (Li and van Staden 1998 ;
Hassan 2006 ; Liu et al. 2006 ; Ohashi et al. 2006 ; Fariduddin et al. 2009 ). Water
or drought stress can stimulate changes in water balance, leaf area expansion,
absorption of photosynthetically active radiation, stomatal closure that reduces
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