Scientia Horticulturae 140 (2012) 66–73
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Scientia Horticulturae
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Antioxidant defense system and proline accumulation enables hot pepper to
perform better under drought
Shakeel Ahmad Anjum
a
, Muhammad Farooq
b,c,∗
, Xio-yu Xie
a
, Xio-jian Liu
a
, Muhammad Furqan Ijaz
d
a
College of Agronomy and Biotechnology, Southwest University, Chongqing 400716, China
b
Department of Agronomy, University of Agriculture, Faisalabad 38040, Pakistan
c
The UWA Institute of Agriculture, The University of Western Australia, Crawley, WA 6009, Australia
d
Institute of Soil and Environmental Sciences, University of Agriculture, Faisalabad 38040, Pakistan
a r t i c l e i n f o
Article history:
Received 13 December 2011
Received in revised form 23 March 2012
Accepted 27 March 2012
Keywords:
Antioxidant enzymes
Drought stress
Lipid peroxidation
Pepper
Proline
a b s t r a c t
Water availability is the most critical factor, which limits the productive potential of plants. Plants adapt to
water deficits by physiological alteration, biochemical changes and osmotic adjustments. The responses
of two pepper (Capsicum annuum L.) cultivars (Shanshu-2001 and Nongchengjiao-2) were investigated to
elucidate the leaf water status, osmolyte accumulation, membrane lipid peroxidation, and key protective
antioxidant enzymes activity under varying levels of progressive drought stress. Pepper cultivars were
submitted to four water supply regimes [80,60, 40 and 20% field capacity (FC)]; served as control, mild,
moderate and severe water stress, respectively. Shanshu-2001 showed a higher relative leaf water con-
tent (RLWC), protein, and proline accumulation than Nongchengjiao-2 in all water regimes during the
course of experiment. Interestingly, total soluble proteins and proline continued to increase with progres-
sion in drought in cultivar Shanshu-2001, whereas in cultivar Nongchengjiao-2 only proline continued to
increase but with a lower rate than cultivar Shanshu-2001. Membrane lipid peroxidation and electrolyte
leakage were increased with prolongation in drought, with higher rates in cultivar Nongchengjiao-2 than
cultivar Shanshu-2001. The progression in drought enhanced the activities of catalase (CAT), peroxidase
(POD) and superoxide dismutase (SOD) initially, which were then decreased. Constitutive activities of
SOD, POD, and CAT were higher in Shanshu-2001 than in Nongchengjiao-2, which resulted in improved
growth and yield in Shanshu-2001. Overall the cultivar Shanshu-2001 was better able to resist drought
as indicated by better growth and yield due to higher antioxidant enzymes, reduced lipid peroxidation,
better accumulation of osmolytes and maintenance of tissue water contents.
© 2012 Elsevier B.V. All rights reserved.
1. Introduction
Changing climate is mainly characterized by increase in emis-
sion of green house gases, global mean temperature, and changes
in precipitation levels and patterns (IPCC, 2007). Owing to changed
patterns of precipitation, episodes of drought are increasing and
are expected to increase in days to come (IPCC, 2007). It is thus
imperative to make the plants ready for the water deficit or
drought. However this all requires better understanding of the
mechanism of resistance and adaptation to drought (Farooq et al.,
2009a).
Plants have evolved complex mechanisms to perceive, respond
and adapt to water deficit. For instance, they can avoid drought
by maximizing water uptake or minimizing water loss (Chaves
∗
Corresponding author at: Department of Agronomy, University of Agriculture,
Faisalabad 38040, Pakistan. Tel.: +92 300 7108652; fax: +92 41 9200605.
E-mail addresses: mfarooq@uaf.edu.pk, farooqcp@gmail.com (M. Farooq).
et al., 2003) or accumulate some osmolytes to cope with the stress
(Szira et al., 2008). These osmolytes accumulate in the cytosol
and are not toxic even at higher concentrations (Reddy et al.,
2004). Drought induces oxidative damage leading in the forma-
tion of active and reactive oxygen species (Farooq et al., 2009a,b).
Production of these species is started with reduction of O
2
lead-
ing in the synthesis of singlet oxygen (
1
O
2
), superoxide (O
2
-
),
hydroxyl radical (OH
-
) or hydrogen peroxide (H
2
O
2
) (Wu et al.,
2008). Production of these species at higher level may damage
cellular membranes and other vital substances like chlorophyll,
DNA, proteins and lipids (Blokhina et al., 2003). Accumulation of
osmolytes, such as proline, glycinebetaine, etc., helps in conserv-
ing tissue water and protect the proteins and cellular membranes
from the osmotic and oxidative stresses (Ashraf and Foolad, 2007;
Farooq et al., 2009a,b). Improvement in the synthesis of these
osmolytes has been found to improve resistance against drought in
maize (Anjum et al., 2011a,b), rice (Farooq et al., 2008a), sunflower
(Hussain et al., 2008); chilling in maize (Farooq et al., 2008b, 2009c),
and heat in sugarcane (Rashid et al., 2011) and wheat (Farooq et al.,
2011).
0304-4238/$ – see front matter © 2012 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.scienta.2012.03.028