INTERNATIONAL JOURNAL OF AGRICULTURE & BIOLOGY

ISSN Print: 15608530; ISSN Online: 18149596

08305/IGC-DYT/2009/111100105
http://www.fspublishers.org

Full Length Article

Drought Stress in Plants: A Review on Morphological

Characteristics and Pigments Composition
CHERUTH ABDUL JALEEL1, PARAMASIVAM MANIVANNAN, ABDUL WAHID, MUHAMMAD FAROOQ, HAMEED
JASIM AL-JUBURI, RAMAMURTHY SOMASUNDARAM AND RAJARAM PANNEERSELVAM
DMJM International (Cansult Maunsell/Aecom Ltd.), Consultant of Gardens Sector Projects, Alain Municipality and Eastern
Emirates, P.O. Box 1419, Al-Ain, Abu Dhabi, United Arab Emirates
Stress Physiology Lab, Department of Botany, Annamalai University, Annamalainagar 608 002, Tamil Nadu, India
Department of Botany, University of Agriculture, Faisalabad-38040, Pakistan
Department of Agronomy, University of Agriculture, Faisalabad-38040, Pakistan
1
Corresponding authors e-mail: abdul79jaleel@yahoo.co.in

ABSTRACT

Plant growth and productivity is adversely affected by natures wrath in the form of various biotic and abiotic stress factors.
Water deficit is one of the major abiotic stresses, which adversely affects crop growth and yield. These changes are mainly
related to altered metabolic functions, one of those is either loss of or reduced synthesis of photosynthetic pigments. This
results in declined light harvesting and generation of reducing powers, which are a source of energy for dark reactions of
photosynthesis. These changes in the amounts of photosynthetic pigments are closely associated to plant biomass yield. This
review describes some aspects of drought induced changes in morphological, physiological and pigments composition in
higher plants.

Key Words: Morphological characters; Secondary metabolism; Pigments composition; Yield

INTRODUCTION of agricultural crops and sustainable food production (Jaleel

et al., 2007b-e; Nakayama et al., 2007). Conventional plant
Stress is an altered physiological condition caused by breeding attempts have changed over to use physiological
factors that tend to disrupt the equilibrium. Strain is any selection criteria since they are time consuming and rely on
physical and chemical change produced by a stress (Gaspar present genetic variability (Zhu, 2002). Tolerance to abiotic
et al., 2002). The term stress is used with various meanings, stresses is very complex, due to the intricate of interactions
the physiological definition and appropriate term as between stress factors and various molecular, biochemical
responses in different situations. The flexibility of normal and physiological phenomena affecting plant growth and
metabolism allows the response initiation to the development (Razmjoo et al., 2008). High yield potential
environmental changes, which fluctuate regularly and are under drought stress is the target of crop breeding. In many
predictable over daily and seasonal cycles. Thus every cases, high yield potential can contribute to yield in
deviation of a factor from its optimum does not necessarily moderate stress environment (Blum, 1996).
result in stress. Stress being a constraint or highly un- Drought stress is considered to be a moderate loss of
predictable fluctuations imposed on regular metabolic water, which leads to stomatal closure and limitation of gas
patterns cause injury, disease or aberrant physiology. Plants exchange. Desiccation is much more extensive loss of
are frequently exposed to many stresses such as drought, water, which can potentially lead to gross disruption of
low temperature, salt, flooding, heat, oxidative stress and metabolism and cell structure and eventually to the
heavy metal toxicity, while growing in nature. cessation of enzyme catalyzed reactions (Smirnoff, 1993;
Drought is a meteorological term and is commonly Jaleel et al., 2007d). Drought stress is characterized by
defined as a period without significant rainfall. Generally reduction of water content, diminished leaf water potential
drought stress occurs when the available water in the soil is and turgor loss, closure of stomata and decrease in cell
reduced and atmospheric conditions cause continuous loss enlargement and growth (Fig. 1). Severe water stress may
of water by transpiration or evaporation. Drought stress result in the arrest of photosynthesis, disturbance of
tolerance is seen in almost all plants but its extent varies metabolism and finally the death of plant (Jaleel et al.,
from species to species and even within species. Water 2008c).
deficit and salt stresses are global issues to ensure survival Water stress inhibits cell enlargement more than cell

To cite this paper: Jaleel, C.A., P. Manivannan, A. Wahid, M. Farooq, R. Somasundaram and R. Panneerselvam, 2009. Drought stress in plants: a review on
morphological characteristics and pigments composition. Int. J. Agric. Biol., 11: 100105
 JALEEL et al. / Int. J. Agric. Biol., Vol. 11, No. 1, 2009

division. It reduces plant growth by affecting various varieties of a species show great differences in the
physiological and biochemical processes, such as production of roots (e.g., rice; Fig. 3). The importance of
photosynthesis, respiration, translocation, ion uptake, root systems in acquiring water has long been recognized. A
carbohydrates, nutrient metabolism and growth promoters prolific root system can confer the advantage to support
(Jaleel et al., 2008a-e; Farooq et al., 2008). In plants, a accelerated plant growth during the early crop growth stage
better understanding of the morpho-anatomical and and extract water from shallow soil layers that is otherwise
physiological basis of changes in water stress resistance easily lost by evaporation in legumes (Johansen et al.,
could be used to select or create new varieties of crops to 1992). The development of root system increases the water
obtain a better productivity under water stress conditions uptake and maintains requisite osmotic pressure through
(Nam et al., 2001; Martinez et al., 2007). The reactions of higher proline levels in Phoenix dactylifera (Djibril et al.,
plants to water stress differ significantly at various 2005). An increased root growth due to water stress was
organizational levels depending upon intensity and duration reported in sunflower (Tahir et al., 2002) and Catharanthus
of stress as well as plant species and its stage of growth roseus (Jaleel et al., 2008a & c). The root dry weight was
(Chaves et al., 2002; Jaleel et al., 2008b). Understanding decreased under mild and severe water stress in Populus
plant responses to drought is of great importance and also a species (Wullschleger et al., 2005). An increase in root to
fundamental part for making the crops stress tolerant shoot ratio under drought conditions was related to ABA
(Reddy et al., 2004; Zhao et al., 2008). content of roots and shoots (Sharp & LeNoble, 2002;
Effects of drought stress on morphological Manivannan et al., 2007b). The root growth was not
characteristics. It has been established that drought stress is significantly reduced under water deficits in maize and
a very important limiting factor at the initial phase of plant wheat (Sacks et al., 1997).
growth and establishment. It affects both elongation and Greater plant fresh and dry weights under water
expansion growth (Anjum et al., 2003a; Bhatt & Srinivasa limited conditions are desirable characters. A common
Rao, 2005; Kusaka et al., 2005; Shao et al., 2008). Among adverse effect of water stress on crop plants is the reduction
the crops, rice as a submerged crop, is probably more in fresh and dry biomass production (Farooq et al., 2009).
susceptible to drought stress than most other plant species Plant productivity under drought stress is strongly related to
(Fig. 2). In soybean, the stem length was decreased under the processes of dry matter partitioning and temporal
water deficit conditions (Specht et al., 2001). The plant biomass distribution (Kage et al., 2004). Diminished
height was reduced up to 25% in water stressed citrus biomass due to water stress was observed in almost all
seedlings (Wu et al., 2008). Stem length was significantly genotypes of sunflower (Tahir & Mehid, 2001). However,
affected under water stress in potato (Heuer & Nadler, some genotypes showed better stress tolerance than the
1995), Abelmoschus esculentus (Sankar et al., 2007 & 08); others. Mild water stress affected the shoot dry weight,
Vigna unguiculata (Manivannan et al., 2007a); soybean while shoot dry weight was greater than root dry weight loss
(Zhang et al., 2004) and parsley (Petroselinum crispum) under severe stress in sugar beet genotypes (Mohammadian
(Petropoulos et al., 2008). et al., 2005). Reduced biomass was seen in water stressed
Water stress greatly suppresses cell expansion and cell soybean (Specht et al., 2001), Poncirus trifoliatae seedlings
growth due to the low turgor pressure. Osmotic regulation (Wu et al., 2008), common bean and green gram (Webber et
can enable the maintenance of cell turgor for survival or to al., 2006) and Petroselinum crispum (Petropoulos et al.,
assist plant growth under severe drought conditions in pearl 2008). A moderate stress tolerance in terms of shoot dry
millet (Shao et al., 2008). The reduction in plant height was mass plants was noticed in rice (Lafitte et al., 2007).
associated with a decline in the cell enlargement and more Yield and related traits. Fetching greater harvestable yield
leaf senescence in A. esculentus under water stress (Bhatt & is the ultimate purpose of growing crops. The crop species
Srinivasa Rao, 2005). Development of optimal leaf area is show great differences for final harvestable yield under
important to photosynthesis and dry matter yield. Water drought stress (Fig. 4). In early plantings of sunflower, the
deficit stress mostly reduced leaf growth and in turn the leaf yield increase was associated with both an increase in grain
areas in many species of plant like Populus (Wullschleger et number and in individual grain weight (Soriano et al.,
al., 2005), soybean (Zhang et al., 2004) and many other 2002). The partitioning of dry matter to the head is critical
species (Farooq et al., 2009). Significant inter-specific in the process of yield determination in water stressed
differences between two sympatric Populus species were parsley (Petropoulos et al., 2008). The effect of water
found in total number of leaves, total leaf area and total leaf deficits on the harvest index of sunflower is complex due to
biomass under drought stress (Wullschleger et al., 2005). the interactions between the timing and intensity of the
The leaf growth was more sensitive to water stress in wheat stress relative to the developmental processes that determine
than in maize (Sacks et al., 1997); Vigna unguiculata the components of yield (Soriano et al., 2002). Exposure of
(Manivannan et al., 2007a) and sunflower (Manivannan et sunflower plants to drought stress at bud initiation stage was
al., 2007b & 2008). more detrimental to seed and biological yield than at seed
Production of ramified root system under drought is filling stage (Prabhudeva et al., 1998). The yield
important to above ground dry mass and the plant species or components like grain number and grain size were

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decreased under pre-anthesis drought stress treatment in Fig. 1. Some important causes of growth reduction in
wheat (Edward & Wright, 2008). In some other studies on plants under drought stress
maize, drought stress greatly reduced the grain yield, which
was dependent on the level of defoliation due to water stress
during early reproductive growth (Kamara et al., 2003;
Monneveux et al., 2006). Water stress reduces seed yield in
soybean usually as a result of fewer pods and seeds per unit
area (Specht et al., 2001).
In water stressed soybean the seed yield was far below
when compared to well-watered control plants (Specht et
al., 2001). Water stress reduced the head diameter, 100-
achene weight and yield per plant in sunflower. There was a
negative correlation of head diameter with fresh root and
shoot weight, while a positive one between dry shoot weight
and achene yield per plant under water stress (Tahir &
Mehid, 2001). Water stress for longer than 12 days at grain
filling and flowering stage of sunflower (grown in sandy
loam soil) was the most damaging in reducing the achene Fig. 2. Effect of drought stress on the vegetative growth
yield in sunflower (Mozaffari et al., 1996; Reddy et al., of rice (cv. IR64). Both the plants were grown under
2004), seed yield in common bean and green gram (Webber well-watered conditions up to 20 days following
et al., 2006), maize (Monneveux et al., 2006) and emergence. One pot (on the right) was submitted to
Petroselinum crispum (Petropoulos et al., 2008). progressive soil drying (drought stress) for 20 days.
Effects of drought stress on pigment composition. The decrease in soil moisture was controlled by partial
Photosynthetic pigments are important to plants mainly for re-watering of the stressed pots to avoid a quicker
harvesting light and production of reducing powers. Both imposition of stress and to homogenize the
the chlorophyll a and b are prone to soil drying (Farooq et development of drought stress. Well-watered control
al., 2009). However, carotenoids have additional roles and pot was maintained at initial target weight by adding
partially help the plants to withstand adversaries of drought. the daily water loss back to the pot
A brief account of changes in chlorophylls and caroteniods
in drought stressed plants is given below:
Chlorophylls. Drought stress produced changes in the ratio
of chlorophyll a and b and carotenoids (Anjum et al.,
2003b; Farooq et al., 2009). A reduction in chlorophyll
content was reported in drought stressed cotton (Massacci et
al., 2008) and Catharanthus roseus (Jaleel et al., 2008a-d).
The chlorophyll content decreased to a significant level at
higher water deficits in sunflower plants (Kiani et al., 2008)
and in Vaccinium myrtillus (Tahkokorpi et al., 2007). The
foliar photosynthetic rate of higher plants is known to
decrease as the relative water content and leaf water
potential decreases (Lawlor & Cornic, 2002). However, the
debate continues as, whether drought mainly limits
photosynthesis through stomatal closure or through
metabolic impairment (Lawson et al., 2003; Anjum et al.,
2003b). Both stomatal and non-stomatal limitation was
generally accepted to be the main determinant of reduced
photosynthesis under drought stress (Farooq et al., 2009).
The limitation of photosynthesis under drought through
metabolic impairment is more complex phenomenon than
stomatal limitation and mainly it is through reduced
photosynthetic pigment contents in sunflower (Reddy et al.,
2004). Chlorophyll b content increased in two lines of okra, Carotenoids. Carotenoids are a large class of isoprenoid
whereas chlorophyll a remained unaffected resulting in a molecules, which are de novo synthesized by all
significant reduction in Chl a: b ratio in both cultivars under photosynthetic and many non-photosynthetic organisms
water limiting regimes (Estill et al., 1991; Ashraf et al., (Andrew et al., 2008). They are divided into the
1994). hydrocarbon carotenes, such as lycopene and -carotene or

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Fig. 3. Root growth and proliferation under well- effects of ROS at this site is essential for chloroplast
watered and drought stress conditions in various rice functioning. Here -carotene, in addition to function as an
genotypes grown in root box. Rice genotypes, Nip, sl 13, accessory pigment, acts as an effective antioxidant and plays
sl 34, sl 45 and sl 50 were grown under well-watered a unique role in protecting photochemical processes and
and drought (15% soil moisture contents) stress and sustaining them (Havaux, 1998). A major protective role of
harvested 38 days after seeding (courtesy Ms. Mana -carotene in photosynthetic tissue may be through direct
Kano) quenching of triplet chlorophyll, which prevents the
generation of singlet oxygen and protects from oxidative
damage (Farooq et al., 2009).
Improving pigments synthesis. Water stress, among other
changes, has the ability to reduce the tissue concentrations
of chlorophylls and carotenoids (Havaux, 1998; Kiani et al.,
2008), primarily with the production of ROS in the
thylakoids (Niyogi, 1999; Reddy et al., 2004). However,
reports dealing with the strategies to improve the pigments
contents under water stress are entirely scarce. The available
reports show that exogenous application of brassinolide,
uniconazole and methyl jasmonate improved the drought
tolerance with increased activities of SOD, CAT and APX,
ABA and total improved carotenoid contents in maize (Li et
al., 1998), while methyl jasmonate brought about a
threefold increase in the -carotene synthesis as well as
degradation of the cholorophyll contents in the epidermal
peels (Prez et al., 1993). Likewise, an important role of
Fig. 4. Loss of harvestable yield in different crop tocopherols, lipid-soluble antioxidant in chloroplasts, has
species when the drought stress was applied at been envisioned in improved pigments contents under stress
reproductive stage (as indicated in the sources conditions in the photosynthetic organisms including
mentioned in the bars) tobacco (Tanaka et al., 1999) and Arabidopsis thaliana and
Synechocystis sp. PCC6803 (DellaPenna & Pogson, 2006).
These data warrant concerted efforts on the either the
induction of pigment synthesis or modification of pigment
biosynthesis pathways for enhanced drought tolerance in
plants.

CONCLUSION

Drought stress affects the growth, dry mater and

harvestable yield in a number of plant species, but the
tolerance of any species to this menace varies remarkably. A
ramified root system has been implicated in the drought
tolerance and high biomass production primarily due to its
ability to extract more water from soil and its transport to
xanthophylls, typified by lutein (Jaleel et al., 2007c). aboveground parts for photosynthesis. In addition to other
Oxidative damage generated by drought stress in the plant factors, changes in photosynthetic pigments are of
tissue is alleviated by a concerted action of both enzymatic paramount importance to drought tolerance. Of the two
and non-enzymatic antioxidant systems. These include - photosynthetic pigments classes, carotenoids show
carotenes, ascorbate (AA), -tocopherol (-toc), reduced multifarious roles in drought tolerance including light
glutathione (GSH) and enzymes including superoxide harvesting and protection from oxidative damage caused by
dismutase (SOD), peroxidase (POD), ascorbate peroxidase drought. Thus, increased contents specifically of carotenoids
(APX), catalase (CAT), polyphenol oxidase (PPO) and are important for stress tolerance.
glutathione reductase (GR; Prochazkova et al., 2001).
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