Introduction
The Intergovernmental Panel on Climate Change (IPCC) assessed the extent to
which recent observed changes in natural biological systems have been caused by
climate change. Most field biologists are convinced that they are already seeing
important biological impacts of climate change14,69; however, they have
encountered difficulty in convincing other academic disciplines, policy-makers and
the general public. Since 1900, concentration of CO 2 has risen from 290 mol mol-1
to 377 mol mol-1 in 2004 (Keeling & Whorf, 2005), and is currently increasing at a
rate of 1.5 (0.9-2.8) mol mol -1 a-1, which is mainly due to fossil fuel combustion
and deforestation (IPCC, 2001). The prediction is that CO 2 concentration will rise to
over 500 mol mol-1 by 2050 and to almost 700 mol mol-1 by 2100, which will
result in an increase in global average temperature of approximately 2.5 3.5 C
(IPCC, 2001) and forest regions at high latitudes can be heated by almost 10 C
(Christensen et al. 2007). Furthermore, since CO
revenue (Mora et al., 2015). However, plant growth is strongly limited by climate
variables such as air temperature, water availability, and solar radiation [1-4], which
are changing in response to ongoing climate change. Terrestrial plants have
evolved complex systems of water management and water conservation that utilize
a suite of adaptive morphological traits (e.g., small vessel elements, thick waxy
cuticles, low specific leaf areas) and physiological processes (e.g., hormonal
signaling, stomatal action), which collectively control plant-water relations and
drought-resistance. Large differences have developed among plant taxa in their
abilities to cope with moisture stress as a result of climate-related natural selection
(Bansal et al., 2015). Bansal et al, 2005 found there was also considerable genetic
variation in drought-resistance among populations originating along climate
gradients. They found that populations originating from warmer, drier climates
generally had lower values of transpiration, water deficit and SLA. Even though
populations from warm climates had relatively high drought-resistance, populations
originating from the coolest climates also exhibited increased drought- resistance.
Photosynthesis
Most plant species only grow in a certain temperature range. Thus, some are likely
to adapt to warmer temperatures by changing their growth and development or by
shifting their ranges, provided that the optimum temperatures are not exceeded.
Some species may fail to adapt to this global change and may even become
extinct if the air temperature is too high. Because the leaf is the key organ
performing photosynthesis and transpiration, its development, which varies with
environmental factors, is an important determinant of total plant productivity
[Thomas et al.,
Air warming and soil drought may have very distinct effects on trees, and their
synergic impact remains unclear (Mittler 2006; Rennenberg et al. 2006).
Temperature can affect photosynthesis through modulation of the rates of activity
of photosynthetic enzymes and the electron transport chain (Sage & Kubien 2007)
and, in a more indirect manner, through leaf temperatures defining the magnitude
of the leaf-to- air vapour pressure difference, a key factor influencing stomatal
conductances. Photosynthetic response to soil drought and air warming is highly
complex. Air warming increases photosynthesis as long as the plant optimal
temperature is not exceeded; after that, photorespiration is stimulated and
photosynthesis inhibited (Berry & Bjo rkman 1980; Brooks & Farquhar 1985; Lea &
Leegood 1999). Moreover, high temperatures can directly affect different phases of
photosynthesis by inhibiting the activity of the both enzimes: ribulose- 1,5carboxylase/oxygenase as well as processes associated with the regeneration of
Rubiscos substrate, ribulose-1,5-bisphosphate (RuBP) through the Calvin cycle
and Rubisco activase enzyme, essential in Rubisco activation, and damage the
thermolabile photosystem II (PSII) (Haldimann & Feller 2004). Stomata open to
maximise dissipation of latent heat (Rennenberg et al. 2006). An increased
assimilation of carbon by plants, can be expected with an elevation of CO 2
concentration and this could increase the amount of carbon stored in vegetation
and below ground (i.e. roots and soil organic matter).
Mn-SOD and Fe-SOD are phylogenetically related to each other and they are very
similar in their primary and tertiary structures, whereas Cu/Zn-SOD shows different
structural features (Fink & Scandalios, 2002). Foyer and Allen (2003) suggest that
redox signalling was the first type of sensory regulation that evolved in nature.
Sequence similarities in higher plants to cyanobacterial redox signalling
components indicate homology and suggest conserved sensory and signalling
functions (Forsberg et al., 2001). Photosynthetic organisms have perfected the art
of redox control. It is now widely accepted that redox signals are key regulators of
plant metabolism, morphology, development, growth, and eventual death (Foyer &
Allen, 2003). One of the major ways in which plants transmit information about the
changing environmental factors is the ROS sensing, producing and scavenging
system. Abiotic and biotic stressors regardless of the first target site of their action,
affect the cellular balance between different redox buffers and oxidants, called
redox homeostasis (Apel & Hirt, 2004; Dizengremel et al. , 2009). A lot of evidence
points to this phenomenon as a common background of most, if not all,
environmental stresses perceived not only as a source of oxidative stress, but also
as a mechanism controlling the main aspects of plant adaptation to various growth
conditions (Foyer & Noctor, 2005a; 2009). The interplay between ROS production
and scavenging determines the steady-state level of ROS in cells, as well as the
ROS signature, i.e. the duration, localization, and amplitude of ROS signals
conditioning stress responses (Mahalingham & Fedoroff, 2003; Miller et al ., 2008).
The ascorbate-glutathione (AA-GSH) cycle serves as the main antioxidant pathway
in plant cells linking the protection against ROS to the redox-regulated plant
acclimation response (Foyer & Noctor, 2005b). The AA-GSH cycle involves
successive oxidations and reductions of ascorbate and glutathione catalysed by
the enzymes constituting the cycle, namely ascorbate peroxidase (APX, EC
1.11.1.11),
monodehydroascorbate
reductase
(MDHAR,
EC
1.6.5.4),
from its direct antioxidant role, the AA-GSH cycle functions in ROS sensing and
signalling (Foyer & Noctor, 2005a, 2005b). Recent genetic evidence suggests that
ROS do not trigger PCD or senescence by causing damage to the cell but they act
as signals that activate pathways of gene expression that lead to genetically
regulated cell death (Foyer & Noctor, 2005a; Dietz, 2008).
Under these stressful conditions imbalance between light energy absorbed through
PSII and the ultimate consumption of the photosynthetic electrons through
metabolic pathways
Pinus
banksiana was less when growing at relatively low temperatures. This was
attributable in part to cold induced production of filtering anthocyanin in the
epidermis and in part because of increased photosynthetic capacity, most likely
caused by an increased capacity at low temperatures to keep QA, the first stable
quinone electron acceptor of PSII, oxidized at high irradiance (Krol et al., 1995).
Antioxidant enzymes also play a role in protecting against freezing
temperatures (Tao et al., 1998; Garca-Plazaola et al., 1999), even in roots (Zhao &
Blumwald, 1998). But if night time temperatures are too low, photoinhibition
increases. Under such conditions a 1C increase in average night-time
temperature allowed Abies lasiocarpa seedlings to avoid photoinhibition during the
day, and increased photosynthesis by 650% (Germino & Smith, 1999). By contrast,
80% shading increased photo-synthesis by only 40%.
Drought
Drought is a globally important abiotic factor limiting tree growth, with acute
drought events commonly causing forest and woodland dieback (Allen et al. 2010).
Forests contain up to 80% of the global above-ground and 40% of the belowground carbon in terrestrial ecosystems (Dixon et al. 1994). Therefore, the impact
of drought on trees might become an important factor influencing the global carbon
cycle and thereby further boost global warming because of reduced CO2 fixation.
Trees have evolved various mechanisms which help them to cope with limited
water supply. Responses to drought include: (1) reducing the water deficit by
developing root systems able to take up water deep in the soil; (2) minimizing
water losses through stomatal closure and producing small leaves; and (3)
accumulating osmoprotective substances (Kozlowski & Pallardy 2002).
Drought may become more severe and more prevalent with predictions of
raising global air temperatures and, locally, decreasing rainfall, especially in areas
where problems with water supply already exist (IPCC 2007). Predictions are that
more areas may become affected by drought and, as a result, forest health, growth
and productivity may be impaired (IPCC 2007). Potential adaptive management
may include the preferred selection of more tolerant species or genotypes in some
areas (Marris 2009). The determination of the degree of drought tolerance of tree
and shrub species, or of genotypes within species, is currently not straightforward,
especially if selections are desired at a seedling or sapling stage. A number of
underlying physiological mechanisms and biochemical compounds have been
proposed as markers for drought stress and tolerance, among them anti-oxidative
and photoprotective defence systems (Smirnoff 1993, Tausz et al. 2004).
Antioxidative and photoprotective defence systems keep reactive oxygen species,
produced under conditions of photo-oxidative stress, under control. Upon drought
exposure, photo-oxidative stress can occur when stomatal closure restricts CO 2
input into the leaf (Smirnoff 1993, Chaves et al. 2009). Reducing equivalents
(NADPH) produced in the light reactions which are not used in carbon fixation can
be consumed in alternative pathways such as Mehler reaction or pho-torespiration,
resulting in the production of ROS (Flexas and Medrano 2002a, Asada 2006). If
consumption of reducing equivalents decreases, the imbalances between electron
use and energy consumption under continuing light-driven electron transport can
cause a situation of 'excess excitation energy' (Flexas and Medrano 2002a, Tausz
et al. 2004, Asada 2006). Excess energy in itself as well as the alternative electron
transport pathways produce ROS. Reactive oxygen species can disrupt cell
membranes, nucleic acids and other cellular functions (Asada 2006, Gill and Tuteja
2010). Despite their potentially destructive consequences, ROS can also act as
signal and sensing factors triggering plant stress responses (Foyer and Noctor
2009). Trees can cope with excess excitation energy and photo-oxidative stress in
several ways. Acclimation to high light conditions can include decreased
chlorophyll contents, which reduces the amount of light absorbed in the first
place (Niinemets 2007). The thylakoid pigment zeaxanthin is involved in the
dissipation of excess energy as heat (Flexas and Medrano 2002a,
Demmig-
Adams and Adams 2006). Dissipation of excess energy can minimize the formation
of ROS and prevent photoinhibition. If energy dissipation is insufficient, ROS
generation in the photosynthetic membranes or as a result of photorespiration or
the Mehler reaction can be enhanced (Noctor et al. 2002, Tausz et al. 2004).
Reactive oxygen species are detoxified by antioxidants, and both antioxidants and
ROS have important role in signalling pathways in response to stress (MunnBosch et al. 2012). Low-molecular-weight antioxidants (carotenoids, ascorbic acid,
glutathione and -tocopherol) function as alternative electron acceptors (Tausz et
al. 2004). These antioxidants are involved in essential and complex pathways,
where excess energy is eliminated and ROS detoxified (Foyer and Shigeoka
2010). While antioxidants can serve as chemical reductants, their functions and
their regeneration are also linked in enzymatic cycles, such as the ascorbateglutathione cycle where ascorbic acid is regenerated by glutathione (Foyer and
Noctor 2011). Ascorbic acid has a role in conversion of zeaxanthin in the
xanthophyll cycle and also in the regeneration of -tocopherol, a membrane-bound
lipophilic antioxidant (Foyer and Shigeoka 2010, Munn-Bosch et al. 2012).
Besides taking part in scavenging of ROS, -tocopherol functions as a stabilizer of
cell membranes (Munn-Bosch and Alegre 2002). Glutathione and ascorbic acid
are near-ubiquitous water-soluble antioxidants and they take part in many
physiological processes (i.e., growth regulation), apart from their defence role
under stress (Foyer and Noc tor 2011, Smirnof f 2011). They are present in two
forms: reduced and oxidized (Apel and Hirt 2004, Foyer and Noctor 2011), which
together make up the total pool of the antioxidant. During the removal of ROS,
antioxidants are transformed into their oxidized form, and must be regenerated
(Apel and Hirt 2004, Foyer and Noctor 2011). Antioxidants are normally thought to
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