Journal of Plant Physiology 169 (2012) 1392–1400
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Journal of Plant Physiology
j ourna l ho mepage: www.elsevier.de/jplph
Growth under elevated atmospheric CO
2
concentration accelerates leaf
senescence in sunflower (Helianthus annuus L.) plants
Lourdes de la Mata, Purificación Cabello, Purificación de la Haba, Eloísa Agüera
∗
Departamento de Botánica, Ecología y Fisiología Vegetal, Área de Fisiología Vegetal, Facultad de Ciencias, Universidad de Córdoba, Campus de Rabanales, Edificio Celestino Mutis
(C4), 3
a
planta, E-14071 Córdoba, Spain
a r t i c l e i n f o
Article history:
Received 5 February 2012
Received in revised form 24 April 2012
Accepted 21 May 2012
Keywords:
Elevated CO2
Hexoses
Oxidative status
Photosynthetic pigments
Senescence
Sunflower
a b s t r a c t
Some morphogenetic and metabolic processes were sensitive to a high atmospheric CO
2
concentration
during sunflower primary leaf ontogeny. Young leaves of sunflower plants growing under elevated CO
2
concentration exhibited increased growth, as reflected by the high specific leaf mass referred to as dry
weight in young leaves (16 days). The content of photosynthetic pigments decreased with leaf develop-
ment, especially in plants grown under elevated CO
2
concentrations, suggesting that high CO
2
accelerates
chlorophyll degradation, and also possibly leaf senescence. Elevated CO
2
concentration increased the
oxidative stress in sunflower plants by increasing H
2
O
2
levels and decreasing activity of antioxidant
enzymes such as catalase and ascorbate peroxidase. The loss of plant defenses probably increases the
concentration of reactive oxygen species in the chloroplast, decreasing the photosynthetic pigment con-
tent as a result. Elevated CO
2
concentration was found to boost photosynthetic CO
2
fixation, especially in
young leaves. High CO
2
also increased the starch and soluble sugar contents (glucose and fructose) and
the C/N ratio during sunflower primary leaf development. At the beginning of senescence, we observed
a strong increase in the hexoses to sucrose ratio that was especially marked at high CO
2
concentration.
These results indicate that elevated CO
2
concentration could promote leaf senescence in sunflower plants
by affecting the soluble sugar levels, the C/N ratio and the oxidative status during leaf ontogeny. It is likely
that systemic signals produced in plants grown with elevated CO
2
, lead to early senescence and a higher
oxidation state of the cells of these plant leaves.
© 2012 Elsevier GmbH. All rights reserved.
Introduction
Continuous emissions of CO
2
from the burning of fossil fuels are
expected to raise global atmospheric CO
2
concentrations. Human
activities not only affect CO
2
concentrations, but also alter the
global nitrogen cycle by increasing the inputs of fixed forms of
nitrogen, mainly through extensive use of chemical fertilizers. The
Intergovernmental Panel on Climate Change (IPCC) has predicted
that the CO
2
concentration may increase by 660–790 L L
-1
from
2060 to 2090 (IPCC, 2007). This is expected to raise global temper-
atures due to the CO
2
capacity to absorb infrared light (Schneider,
1989; Taylor and MacCracken, 1990). Therefore, continuous emis-
sions of this gas at high levels are believed to cause climate change.
One of the most obvious effects of climate change is its effect
Abbreviations: APX, ascorbate peroxidase; DW, dry weight; ROS, reactive oxygen
species; RuBP, ribulose-1,5-bisphosphate; rubisco, ribulose-1,5-bisphophate car-
boxylase/oxygenase; SLM, specific leaf mass; XET, xyloglucan endotransglycosidase.
∗
Corresponding author. Tel.: +34 957218367; fax: +34 957211069.
E-mail addresses: b42matsl@uco.es (L. de la Mata), bv1cahap@uco.es (P. Cabello),
bv1hahep@uco.es (P. de la Haba), vg1agbue@uco.es (E. Agüera).
on living beings, especially on plants, which have been found to
exhibit alterations potentially affecting some steps of their growth
cycle. Studies on various plant species have suggested that climate
changes will affect the development, growth and productivity of
plants through alterations in their biochemical, physiological and
morphogenetic processes (Bazzaz and Fajer, 1992).
Senescence is a stage of the plant growth cycle that involves
strong metabolic and structural changes. Markers associated with
leaf senescence in sunflower plants have shown that senescence
initiates and progresses in primary leaves aged between 28 and
42 days (Cabello et al., 2006). Senescence typically involves ces-
sation of photosynthesis and degeneration of cellular structures,
with strong losses of chlorophyll (Ougham et al., 2008), carotenoids
and proteins and a great increase of lipid peroxidation (Srivalli
and Khanna-Chopra, 2004; Agüera et al., 2010). Senescence is not
only a degenerative process, but also a recycling process by which
nutrients are translocated from senescing cells to young leaves,
developing seeds or storage tissues (Gan and Amasino, 1997).
Leaf senescence is therefore an active, highly regulated and pro-
grammed degeneration process, required for plant survival and
controlled by multiple developmental and environmental signals
(Lim et al., 2003). Senescence induction and development are both
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http://dx.doi.org/10.1016/j.jplph.2012.05.024