Scientia Horticulturae 217 (2017) 164–172
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Scientia Horticulturae
journal homepage: www.elsevier.com/locate/scihorti
The physiological responses of various pomegranate cultivars to
drought stress and recovery in order to screen for drought tolerance
Mohammadreza Pourghayoumi
a
, Davood Bakhshi
a,∗
, Majid Rahemi
b
,
Ali Akbar Kamgar-Haghighi
c
, Ali Aalami
d
a
Department of Horticultural Sciences, Faculty of Agricultural Sciences, University of Guilan, Rasht, Iran
b
Department of Horticultural Sciences, Faculty of Agricultural Sciences, Shiraz University, Shiraz, Iran
c
Department of Irrigation, Faculty of Agricultural Sciences, Shiraz University, Shiraz, Iran
d
Department of Plant Biotechnology, Faculty of Agricultural Sciences, University of Guilan, Rasht, Iran
a r t i c l e i n f o
Article history:
Received 17 August 2016
Received in revised form 24 January 2017
Accepted 26 January 2017
Keywords:
Punica granatum
Water stress
Photosynthesis
Recovery
Tolerant
a b s t r a c t
In order to screen pomegranate cultivars for drought tolerance, few rapid, less expensive and reliable
methods were used. Two-year-old pomegranate (Punica granatum L.) plants of various commercial culti-
vars namely Rabab-e-Neyriz’ (‘Rabab’), ‘Shishe-cap-e-Ferdows’ (‘Shishecap’), ‘Malas-e-Saveh’ (M-Saveh),
‘Malas-e-Yazdi’ (‘M-Yazdi’), and ‘Ghojagh-e-Qom’ (‘Ghojagh’) were grown in large containers filled with
a mixture of leaf mould, sand, and soil (1:1:1, by volume) in greenhouse. The plants were subjected to
14-day drought stress by withholding irrigation, followed by re-watering for 7 days. Midday stem water
potential (
stem
), leaf relative water content (RWC), membrane stability index (MSI), leaf dry mass per
area (LMA), rapid test for drought tolerance (DTI), gas exchange parameters including net photosynthe-
sis (A
n
), leaf scale transpiration (T
r
), and stomatal conductance (g
s
), and intrinsic water use efficiency
(IWUE) were determined in well-watered and drought-stressed plants. All cultivars showed an ability
to tolerate drought stress, but ‘Ghojagh’ exhibited more tolerance, with a higher RWC and
stem
and
a greater osmotic adjustment. ‘Ghojagh’ also revealed higher cell membrane stability and IWUE and a
lower reduction in net CO
2
assimilation rate. This study found that ‘M-Yazdi’ was more vulnerable to
severe water stress, and displayed the lowest degree of cell membrane stability as compared to the other
examined cultivars and showed no recovery for RWC at the end of recovery period.
© 2017 Elsevier B.V. All rights reserved.
1. Introduction
Pomegranate (Punica granatum L.), from the family Punicaceae,
is a popular fruit of tropical and subtropical regions that is native
to the area stretching from Iran to the Himalayas in northern India
(Fawole and Opara, 2013; Parvizi et al., 2016). It has been cultivated
since ancient times and is mentioned in the Christian bibles and
holy Quran (Rahimi et al., 2012). Drought is the most abiotic stress
factor limiting plant growth and crop production in the world.
Increase in fruit splitting or cracking and decrease in vegetative
growth and yield is economically important losses in pomegranate
orchards resulting from water stress. Drought tolerance is observed
in almost all plant species, but its extent varies from species to
species and even in cultivars of the same species (Jain et al., 2011).
∗
Corresponding author.
E-mail addresses: bakhshi-d@guilan.ac.ir, bakhshidavood@hotmail.com
(D. Bakhshi).
Breeders look for new sources of variations when attempting to find
drought tolerant plants with optimal characteristics for high rates
of photosynthesis and productivity, but tolerance to water stress
is very complex, due to the intricate of the interactions between
stress factors and various physiological, biochemical, and molec-
ular response affecting plant growth (Jaleel et al., 2009; Giancarla
et al., 2012). A better understanding of how drought stress affects
the physiology of the plant helps in the creation and selection of
new drought tolerant cultivars. Plants experience drought stress,
either when the water supply to roots becomes difficult or when
the transpiration rate becomes higher than the rate of water uptake.
These two conditions often occur under arid and semi-arid areas
(Gholami et al., 2012). Different levels of drought stress could be
observed in a plant. A moderate loss of water often leads to stoma-
tal reactions, limiting CO
2
assimilation, stomatal conductance, and
transpiration. Severe water stress, on the other hand, occurs when
most of the protoplasmic water is lost and often leads to a major
disruption of the metabolism, enzymatic antioxidant activities and
http://dx.doi.org/10.1016/j.scienta.2017.01.044
0304-4238/© 2017 Elsevier B.V. All rights reserved.