DEPARTMENT OF PLANT SCIENCE AND BIOTECHNOLOGY

FEDERAL UNIVERSITY OF LAFIA

PSB 111: GROWING PLANTS IN DROUGHT AND SALINE ENVIRONMENT
TOPIC: SALINITY AND DROUGHT STRESS IN PLANTS

BY DR GBENGA AKOMOLAFE

INTRODUCTION

Salinity and drought are two important abiotic stresses that limit the production of food crops
worldwide. Plant responses to salinity and drought are often similar. For example, the first
phase of salinity stress, the osmotic effect, is quite similar to that of drought stress. Plant roots
have effective mechanisms to sense low water potential that result either from an actual lack
of water in the environment, due to low precipitation, or to the presence of excess salt ions in
the water present. In both cases plants cannot uptake sufficient water for normal growth and
development and common stress related signal transduction pathways are activated, for
example increased levels of ABA. Immediately following the induction of these stress related
signal transduction pathways, under drought or salinity, shoot growth, especially leaf growth,
is greatly reduced. A significant contributor to this growth reduction is associated with a change
in cellular oxygen and carbon dioxide levels due to the partial closure of stomata. Plant cells
also produce various reactive oxygen species (ROS) in response to both salt and drought stress.
Increased ROS levels are found in both the apoplast and the cytoplasm. Elevated ROS levels
under drought and salinity result in changes in normal cellular metabolism and cellular redox
homeostasis, and also ROS dependent modification to plant cell-wall structure, which either
restricts cell expansion by cell-wall strengthening, or favours cell expansion by loosening the
wall structure.

Due to climate change, it is predicted that drought and salinity will became more severe in the
upcoming years which could lead to a significant reduction of plant growth and yield of several
economically important species. It has been estimated that worldwide food demand will
increase by 70% until the end of 2050 due to a population growth of 2.3 billion people. In this
context, developing crop plants with high yield and better tolerance to harsh environmental
conditions becomes an urgent need to meet future food demand for next generations. In general,
plant responses to salinity and drought may vary in morphological, physiological and
biochemical aspects and processes. Most of the effects induced by salinity and drought are
negative, however to some extent they can have positive effects as well. It has been reported
that salinity at certain concentrations enhanced plant fecundity due to an increase in
reproduction, but it has also been observed that this enhancement was highly dependent on
genotype and plant developmental stage. Soil water salinity can also have a positive effect on
fine particles helping them to bind together into aggregates, thus improving soil aeration, root
penetration and root growth. Nevertheless, salinity cannot be increased in favor of soil structure
without considering the potential impacts on plant health. Salt-stress resistance represents the
ability of a plant to prevent, reduce or overcome the possible damaging effects caused directly
or indirectly by the presence of excessive soluble salts (accumulation of toxic ions) in its root
zone. A 50% reduction in yield can be considered a measure of salt stress.

Drought stress occurs after a relatively long period with no rains, inducing moisture stress in
the soil detrimental to crop growth, especially in rainfed agriculture. The severity of drought is
strongly related to the timing (growth stage of the plants) and intensity (duration of no rain
period). Other factors such as soil characteristics and agricultural practices can interfere with
crop yields. Previous reports suggest that a positive transgenerational impact on seedling vigor
of Brassica napus has been observed due to drought stress. This phenomenon was explained as
a result of the heterotic effects, altered reservoir of seed storage metabolites, and inter-
generational stress memory formed by stress-induced changes in the epigenome of the seedling.
Compared to salt stress, drought stress has more severe effects on plants and economy but plant
responses are closely related and their defense mechanisms even overlap. The ability of a crop
variety to perform better over other varieties under drought conditions is known as drought
resistance which is linked to achieved yields and potential yields achievable in a given
environment in the absence of drought conditions. Drought resistance is highly environment
specific and yield stability might be influenced by crop management practices, and/or
physiological mechanisms and might not necessarily be associated with the drought resistance
ability of a genotype. In a drought resistant variety, plant growth and development are well-
matched to specific drought environment(s). When sensing salinity or drought stresses, plants
have the capability to combine a range of responses in order to avoid stress injuries and
complete their life cycle. By the activation of various defense mechanisms plants can store
reserves in their organs and use them later for yield production or, they can tolerate stress
conditions without tissue dehydration. Plant-associated organisms play an important role in
improving the adaptation strategies of plants to environmental stresses. In this context,
microorganisms, for example, can rescue plants from the deleterious effects of drought and
salinity through their activity, such as nutrient solubilization, IST and production of
phytohormones (IAA, Cytokinin, ABA or GA), EPS and ACC deaminase. The inoculation of
plants with arbuscular mycorrhizal fungus can also increase plants’ tolerance to short term
salinity exposures.

PHYSIOLOGICAL AND BIOCHEMICAL MECHANISMS OF TOLERANCE OF

PLANTS

Plants develop various physiological and biochemical mechanisms in order to survive in soils

with high salt concentration. Principle mechanisms include, but are not limited to, (1) ion

homeostasis and compartmentalization, (2) ion transport and uptake, (3) biosynthesis of

osmoprotectants and compatible solutes, (4) activation of antioxidant enzyme and synthesis of

antioxidant compounds, (5) synthesis of polyamines, (6) generation of nitric oxide (NO), and

(7) hormone modulation. Some of these mechanisms are discussed below.

Ion Homeostasis and Salt Tolerance

Many plants have developed an efficient method to keep the ion concentration in the cytoplasm

in a low level. Membranes along with their associated components play an integral role in

maintaining ion concentration within the cytosol during the period of stress by regulating ion

uptake and transport. The transport phenomenon is carried out by different carrier proteins,
channel proteins, antiporters and symporters. Maintaining cellular Na+/K+ homeostasis is

pivotal for plant survival in saline environments.

Compatible Solute Accumulation and Osmotic Protection

Compatible solutes, also known as compatible osmolytes, are a group of chemically diverse

organic compounds that are uncharged, polar, and soluble in nature and do not interfere with

the cellular metabolism even at high concentration. They mainly include proline, glycine

betaine, sugar, and polyols. Organic osmolytes are synthesised and accumulated in varying

amounts amongst different plant species. As their accumulation is proportional to the external

osmolarity, the major functions of these osmolytes are to protect the structure and to maintain

osmotic balance within the cell via continuous water influx.

Accumulation of polyols, either straight-chain metabolites such as mannitol and sorbitol or

cyclic polyols such as myo-inositol and its methylated derivatives, is correlated with tolerance

to drought and/or salinity, based on polyol distribution in many species.

Antioxidant Regulation

Abiotic and biotic stress in living organisms, including plants, can cause overflow,

deregulation, or even disruption of electron transport chains (ETC) in chloroplasts and

mitochondria. Under these conditions molecular oxygen (O2) acts as an electron acceptor,

giving rise to the accumulation of ROS. Singlet oxygen (1O2), the hydroxyl radical (OH−), the

superoxide radical (O− 2), and hydrogen peroxide (H2O2) are all strongly oxidizing compounds

and therefore potentially harmful for cell integrity. Antioxidant metabolism, including

antioxidant enzymes and nonenzymatic compounds, play critical parts in detoxifying ROS

induced by salinity stress.

Roles of Polyamines in Salinity Tolerance

Polyamines (PA) are small, low molecular weight, ubiquitous, polycationic aliphatic molecules

widely distributed throughout the plant kingdom. Polyamines play a variety of roles in normal

growth and development such as regulation of cell proliferation, somatic embryogenesis,

differentiation and morphogenesis, dormancy breaking of tubers and seed germination,

development of flowers and fruit, and senescence. It also plays a crucial role in abiotic stress

tolerance including salinity and increases in the level of polyamines are correlated with stress

tolerance in plants.

Roles of Nitric Oxide in Salinity Tolerance

Nitric oxide (NO) is a small volatile gaseous molecule, which is involved in the regulation of

various plant growth and developmental processes, such as root growth, respiration, stomata

closure, flowering, cell death, seed germination and stress responses, as well as a stress

signalling molecule. NO directly or indirectly triggers expression of many redox-regulated

genes. NO reacts with lipid radicals thus preventing lipid oxidation, exerting a protective effect

by scavenging superoxide radical and formation of peroxynitrite that can be neutralised by

other cellular processes. It also helps in the activation of antioxidant enzymes (SOD, CAT,

GPX, APX, and GR). Exogenous NO application has been found to play roles in stress

mitigation, but the effects depend on NO concentration.

Hormone Regulation of Salinity Tolerance

ABA is an important phytohormone whose application to plant ameliorates the effect of stress

condition(s). It has long been recognized as a hormone which is upregulated due to soil water

deficit around the root. Salinity stress causes osmotic stress and water deficit, increasing the

production of ABA in shoots and roots. The accumulation of ABA can mitigate the inhibitory
effect of salinity on photynthesis, growth, and translocation of assimilates. The positive

relationship between ABA accumulation and salinity tolerance has been at least partially

attributed to the accumulation of K+, Ca2+ and compatible solutes, such as proline and sugars,

in vacuoles of roots, which counteract with the uptake of Na+ and Cl− . ABA is a vital cellular

signal that modulates the expression of a number of salt and water deficit-responsive genes.