Plant Adaptations To Salt and Water Stress Dos

Plant Adaptations to Salt and Water Stress: Differences and
Commonalities
RANA MUNNS1
CSIRO Plant Industry, Canberra, and School of Plant Biology,

The University of Western Australia, Perth, Australia
I. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
II. Whole Plant Responses to Water and Salt Stress: Commonalities
and Differences. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
III. Mechanisms of Drought and Salinity Tolerance. . . . . . . . . . . . . . . . . . . . . . . . . . . 6
A. Commonalities ................................................................. 6
B. Long-distance Signals Controlling Leaf Expansion and
Stomatal Conductance........................................................ 7
C. Changes in Leaf Morphology and Root System Architecture.......... 8
D. Turgor Maintenance and Osmotic Adjustment........................... 9
E. Differences—Salt Specific .................................................... 9
IV. Adaptations, Traits and QTLs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
A. Drought ......................................................................... 12
B. Salinity .......................................................................... 14
C. Salt-Specific traits.............................................................. 14
D. Traits in Common with Drought ........................................... 19
E. QTL Analysis and Gene Discovery......................................... 20
V. Growth Studies and Experimental Design . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
A. Water Stress .................................................................... 20
B. Salinity .......................................................................... 22
C. Experiments to Distinguish Water Stress from Salt-Specific Effects .. 22
1
Corresponding author: E-mail: rana.munns@csiro.au
Advances in Botanical Research, Vol. 57 0065-2296/11 $35.00

Copyright 2011, Elsevier Ltd. All rights reserved. DOI: 10.1016/B978-0-12-387692-8.00001-1
2 RANA MUNNS
VI. Phenotyping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
VII. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
ABSTRACT
Drought and salinity are the most significant abiotic stresses to limit the production of
the world’s staple food crops. Knowledge about physiological traits, and new molec-
ular tools to identify key genes or to provide molecular markers, has the potential to
increase yield over the present limits. The employment of molecular knowledge needs
appropriate experimental design and accurate phenotyping. Testing the value of
physiological traits and key candidate genes is crucial for progress towards crop
improvement on dry and saline land. This chapter reviews specific traits for drought
and salinity tolerance, and experimental methods that could distinguish drought and
salinity adaptations. The use of new phenomics techniques combined with rapidly
advancing molecular tools provides a powerful impetus to identify key traits and
genes for stress tolerance, and new methods to introduce these genes into important
food crops.
I. INTRODUCTION
Water and salt stress due to drought and soil salinity are the two most
intractable abiotic stresses that limit the production of the world’s staple
food crops, wheat and rice. Drought cannot be avoided, and salinity can only
be temporarily reduced.
Scarcity of water will increase in the near future in many regions of the
world due to climate change (IPCC, 2007) and also to urbanization. World
population is predicted to double in the next 50 years, so greater yields must
be extracted from the current agricultural areas along with more marginal
areas (Chaves and Davies, 2010). The need for more water for cities as the
population grows means less is available for irrigation, so a greater propor-
tion of the world’s food will come from drought-prone areas. Productivity of
crops on the best land in the most favourable environment is approaching a
theoretical maximum, so to meet the increasing world food requirement,
productivity in less favourable environments is needed (Passioura and
Angus, 2010).
Salinity is also increasing, due to either salts naturally present in the soil, or
irrigation or the clearing of land for dryland agriculture.
Over 6% of land throughout the world are salt affected, either by salinity
or by the associated condition of sodicity (Munns, 2005). Most of this
salinity, and all of the sodicity, is natural. However, a significant proportion
of recently cultivated agricultural land has become saline due to land clearing
or irrigation. For example, in Australia, about 2 million ha of the 17 million
PLANT ADAPTATIONS TO SALT AND WATER STRESS 3
ha of land farmed by dryland agriculture are affected by secondary salinity to

varying degrees and a further 8 million ha are at risk of salinisation in the
next 40 years (Munns and James, 2003). A report by FAO (reported by
Munns, 2003) stated that of the 230 million ha of irrigated land worldwide
(much of which is in Asia where rice is the main cereal crop), 45 million ha are
salt affected (20%). Irrigated land is only 15% of total cultivated land, but as
irrigated land has at least twice the productivity of rainfed land, it produces
one-third of the world’s food. In Pakistan alone, 1 million ha of the rice-
growing area is affected by salt (Ul Haq et al., 2010).
Natural or ‘primary salinity’ is more widespread than first realised
(Rengasamy, 2006, 2010a), while ‘secondary salinity’, due to land clearing
or irrigation, continues to grow. Hence, increased salt tolerance of crops and
horticultural species is needed to sustain increases in food production in
many regions in the world. Increased salt tolerance of perennial species
used for fodder or fuel production is a key component in reducing the spread
of secondary salinity, while increased salt tolerance of crops will directly
improve production in soils with primary salinity.
Advances in crop productivity come from better management of the land
and from new varieties. Genetic engineering is one way of increasing genetic
gain, but a single gene may have limited effect. There are tens of thousands of
accessions in international seed banks providing a huge genetic diversity, but
selecting is a daunting task. Understanding the importance of various traits,
and the molecular basis will provide a focused approach. This review sum-
marises the physiological understanding of mechanisms of adaptation and
the traits that confer it, and raises the question of how best to screen for these
traits. It also points out the strong commonality between adaptation to water
and salt stress.
Conventional plant breeding with good agronomic practices has doubled
the production of crops in regions that are not limited by water. For example,
the yield of wheat in UK has doubled from 4 to 8 tonnes per hectare over the
past 50 years, but average yield in Australia has increased only from about 1
to 1.5 tonnes per hectare (Richards et al., 2010). This has come about by
empirical selection for yield, which has proved extremely effective, but shows
signs of reaching a plateau. To complement empirical breeding for yield in
water-limited environments, physiological breeding including the use of
molecular markers has been proposed (Cattivelli et al., 2008; Richards
et al., 2002). Knowledge about traits and their mechanistic basis at both
the physiological and molecular level has the potential to take yield improve-
ment further. It is therefore important to recognise and understand the
processes that allow plants to adapt to water and salinity stress, and to
allow an increase in biomass or seed yield for food production.
4 RANA MUNNS
With the knowledge about physiological traits, new molecular tools can be
employed to identify key genes or to provide molecular markers. To do this,
appropriate and accurate phenotyping is needed. Experimental methods that
could distinguish specific traits for drought and salinity adaptations, and
quantify the effect of these traits, will allow QTLs (quantitative trait loci) to
be identified, and from this, the possibility of identification of candidate
genes or least linked molecular markers.
Previous reviews have covered various aspects of the similarities and con-
trasts in the plant response to drought and salt stress, from the whole plant
(Munns, 2002) to the metabolic (Bartels and Sunkar, 2005) and molecular
level (Pardo, 2010), and the use of physiological traits in breeding (Munns and
Richards, 2007). This review focuses on measurable traits for drought and
salinity tolerance, and methods that can be used to select new germ plasm for
improving the growth and yield of crops in dry and saline soil.
II. WHOLE PLANT RESPONSES TO WATER AND SALT

STRESS: COMMONALITIES AND DIFFERENCES
Soil salinity like drought reduces the soil water potential and the ability of
plants to take up water, and this quickly reduces the rate of cell expansion in
growing tissues. The slower formation of photosynthetic leaf area in turns
reduces the flow of assimilates to the meristematic and growing tissues of the
plant, both leaves and roots, although leaves are often more affected than
roots (Munns and Sharp, 1993). At the same time, both water and salt stress
reduce stomatal conductance in the older leaves, which limits their photosyn-
thetic rate. It is difficult to gauge whether this causes an additional reduction
in growth rate, or the reduction in growth rate is in balance with the reduction
in photosynthesis rate, as carbohydrate status of the growing tissue is usually
not affected and sometimes is higher indicating unused assimilate.
With time, salt may exert an additional effect on growth. If excessive
amounts of Naþ or Cl enter the plant it may rise to toxic levels in the
older transpiring leaves. This injury, added to an already reduced leaf area,
will then further limit the flow of carbon compounds to meristems and
growing zones in leaves. The plant may die before seed is produced.
The rate of leaf death is crucial for the survival of the plant. If new leaves
are continually produced at a rate greater than that at which old leaves die,
then there are enough photosynthesising leaves for the plant to produce
flowers and seeds, although in reduced numbers. However, if old leaves die
faster than new ones develop, then the plant may not survive to produce seed.
For an annual plant, there is a race against time to initiate flowers and form
seeds, while the leaf area is still adequate to supply the necessary photosyn-
thate. For perennial species, there is an opportunity to enter a state akin to
dormancy, and thus survive the stress.
The effects of the different components of salinity stress on plant growth
can be visually distinguished. The water stress affects the growth rate of the
younger leaves, the toxic effect causes premature death of older leaves and
can be seen by premature yellowing. The two effects can be quantitatively
measured over time. In a laboratory experiment, when salt is introduced to a
constant level, a two-phase effect on plant growth is evident (Fig. 1). The first
phase of the growth response results from the effect of salt outside the plant.
The salt in the soil solution reduces leaf growth and to a lesser extent root
growth (Munns, 1993). The cellular and metabolic processes involved are in
common to drought-affected plants. Neither Naþ nor Cl builds up in the
growing tissues at concentrations that inhibit growth: meristematic tissues
are fed largely in the phloem from which salt is effectively excluded, and
rapidly elongating cells can accommodate the salt that arrives in the xylem
within their expanding vacuoles.
The second phase of the growth response results from the toxic effect of
salt inside the plant. The salt taken up by the plant concentrates in the old
4
Control
Osmotic effect
Total dry weight (g)
1
Salt Salt-
specific
0 effect
0 10 20 30 40
Time after NaCl added (d)
Fig 1. Growth of two accessions of the diploid wheat progenitor Triticum tauschii
in control solution (closed symbols) and in 150 mM NaCl with supplemental Ca2þ
(open symbols). Circles denote the tolerant accession, triangles the sensitive one. The
arrow marks the time at which symptoms of salt injury could be seen on the sensitive
accession; at that time the proportion of dead leaves was 10% for the sensitive and 1%
for the tolerant accession. Adapted from Munns et al., 1995.
6 RANA MUNNS
A B
100 100
Relative dry matter (%)
80 80
60 60
40 40
20 7.0 dS/m NaCl 20 30.0 dS/m NaCl

7.0 dS/m Hoagland 30.0 dS/m Hoagland
Non-saline Non-saline
0 0
0 5 10 15 20 25 30 35 40 45 0 5 10 15 20 25 30 35 40 45
Number of days of growth
Fig. 2. Cumulative dry matter of wheat (relative to final dry matter in the control
treatment) during 40 days of growth when treated with NaCl or Hoagland nutrient
solution or non-saline (control) at two levels of soil solution EC: (A) 7.0 dS/m and (B)
30.0 dS/m. Reproduced with permission from Rengasamy (2010a); doi 10.1071/
SR09083
leaves: continued transport into transpiring leaves over a long period of time
eventually results in very high Naþ and Cl concentrations, and they die. The
cause of the injury is probably the salt load exceeding the ability of the cells to
compartmentalise salts in the vacuole.
This distinction was illustrated by contrasting genotypes in Fig. 1. Another
way of showing it, in the absence of contrasting genotypes, is to compare
different osmotica with and without NaCl. Non-ionic compounds such as
polyethylene glycol (PEG) are unsuitable for longer term experiments, as it is
taken up over time (reviewed in Munns et al., 2010). The least toxic osmotica is a
balance salt concentration, with the macronutrients in Hoagland’s solution
(Rengasamy, 2010b; Termaat and Munns, 1986). Figure 2 shows the difference
in wheat growth between NaCl versus Hoagland solution at 7 and 30 dS/m.
III. MECHANISMS OF DROUGHT AND

SALINITY TOLERANCE
A. COMMONALITIES
Mechanisms controlling leaf and root growth are in common to drought and
salinity; they are due to factors associated with water stress. This is supported
by the evidence that Naþ and Cl are below toxic concentrations in the
growing cells themselves, in leaves (Fricke, 2004; Hu and Schmidhalter,
1998) and roots (Jeschke, 1984, Jeschke et al., 1986).
The understanding of plant responses to water deficits from the physiolog-

ical to the molecular level has been summarised by Chaves et al. (2003, 2009).
Many of the traits that explain plant adaptation to drought—such as phe-
nology, root size and depth, hydraulic conductivity and the storage of
reserves—are associated with plant development and structure and are con-
stitutive rather than stress induced (Chaves et al., 2003). But a large part of
plant resistance to drought is the ability to dissipate excess radiation, a
concomitant stress under natural conditions. The effects of drought and
salt stresses on photosynthesis are either direct (as the diffusion limitations
through the stomata and the mesophyll and the alterations in photosynthetic
metabolism) or secondary, such as the oxidative stress arising from the
superimposition of multiple stresses (Chaves et al., 2009). The carbon bal-
ance of a plant during a period of salt/water stress and recovery may depend
as much on the velocity and degree of photosynthetic recovery, as it depends
on the degree and velocity of photosynthesis decline during water depletion.
B. LONG-DISTANCE SIGNALS CONTROLLING LEAF EXPANSION AND

STOMATAL CONDUCTANCE
Whether water status, hormonal regulation or supply of photosynthate

exerts dominant control over growth of plants in dry or saline soil is an
issue that has been hotly debated. Over the time scale of days, there is much
evidence to suggest that hormonal signals rather than water relations are
controlling growth in saline soils. The evidence for this is that leaf expansion
in saline soil at the time scale of days does not respond to an increase in leaf
water status (summarised in Munns, 2002). These experiments indicated that
there are chemical signals coming from roots in dry or saline soil that reduce
leaf growth. These are commonly referred to as ‘root signals’. Abscisic acid
(ABA) is the obvious candidate for this signal, as it is found in xylem sap, and
increases after drought and salinity stress (reviewed by Munns and Cramer,
1996). However, there is still no conclusive proof that ABA is the only signal
from the roots; other hormones are involved (reviewed by Dodd, 2005; Pérez-
Alfocea et al., 2010; Ghanem et al., 2011). Moreover, the origin of the ABA
in the xylem sap is not known, for it moves readily in the phloem and
recirculates from leaves to roots (reviewed by Munns and Cramer, 1996).
When roots sense a soil water deficit, root cells change in growth rate and
differentiation, and the root system architecture changes in the degree of
branching or rate of branch root elongation. They also transmit a signal to
the shoot. Although root elongation rate decreases, the decrease is less than
of shoot growth. In Arabidopsis, the proportion of roots as a fraction of
whole plant biomass can increase significantly in drying soil, by at least 30%
8 RANA MUNNS
(Hummel et al., 2010). In grapevines in the field, roots in drying soil continue
to grow into deeper wetter layers, whereas the roots of irrigated plants
proliferate in the topsoil (Lovisolo et al., 2010).
Analysis of Arabidopsis of the impact of water deficit on the activity on
carbon assimilation and of 30 enzymes central to carbon metabolism showed
that drought stress decreased rosette expansion more than photosynthesis,
while root growth was promoted. All C metabolites showed a diurnal turn-
over that increased under water deficit, but with no fundamental reprogram-
ming (Hummel et al., 2010), indicating quantitative shifts in C metabolism
but no induction or repression of novel genes.
Grapevine genotypes differ in their responses to water stress, exhibiting
either isohydric or anisohydric behaviour, which is linked to stomatal and
non-stomatal effects. Stomatal regulation of grapevine is under ABA and
hydraulic control, the latter linked to embolism formation and recovery.
Xylem embolism occurs and repairs during diurnal cycles under ABA con-
trol. Aquaporins are essential mediators of radial transport of water both
across the roots into the xylem, and out from the xylem in leaves. They may
explain the differences in iso- and anisohydric behaviours. Stomatal sensitiv-
ity has genotypic variation in grapevine (Beis and Patakas, 2010), and can be
altered by irrigation practices (Collins et al. 2010).
When roots sense a saline soil, the responses are the same, except that
embolisms do not occur. ABA is not the only signal, at least in species other
than grapevine, as cytokinins, auxins and the ethylene precursor ACC are
implicated (Pérez-Alfocea et al., 2010). Hormonal signals may coordinate
assimilate production and use in the competing sinks of roots, leaves and
lateral shoots. The hormonal regulation of source–sink relations during the
osmotic phase of salinity, the phase when growth rate and development is
reduced and before ions build up to toxic levels in leaves, affects whole plant
energy balance, and is critical to delay the accumulation of ions to toxic levels
(Pérez-Alfocea et al., 2010). Productivity in saline as well as dry soil depends
on maintaining a high growth rate of young leaves, while at the same time
delaying the senescence of older leaves.
C. CHANGES IN LEAF MORPHOLOGY AND ROOT SYSTEM ARCHITECTURE
Hormonal control of cell division and differentiation is clear from the

appearance of leaves, which are smaller in area but often thicker, indicating
that cell size and shape have changed (James et al., 2002). Leaves from salt-
treated plants have a higher weight:area ratio, which means that their tran-
spiration efficiency is higher (more carbon fixed per water lost), a feature that
is common in plants adapted to dry and saline soil.
Recently, the concept of phenotypic plasticity has re-emerged, partly

because of a clearer understanding of the influence of epigenetic regulations
on gene expression. Phenotypic plasticity illustrates the range of phenotypes
that an organism can express as a function of its environment (Schlichting,
1986), that is the resultant component of the G E (genotype by environ-
ment) interaction. Plasticity is more readily seen in morphological traits such
as variation in leaf size or stem length in response to environmental influ-
ences such as shade or herbivory, but can also be measured in complex
physiological traits such as water use efficiency (Nicotra and Davidson,
2010). The genetics and molecular biology underlying such adaptive pheno-
typic plasticity are exciting areas for future research.
Hormonal control of cell division and elongation is also evident in roots.
Several studies have shown that salinity has differential effects on root
elongation rates and lateral root initiation that is due to the osmotic effect
of the saline soil (Munns and Cramer, 1996; Rahnama et al., 2011).
D. TURGOR MAINTENANCE AND OSMOTIC ADJUSTMENT
Turgor maintenance is essential for cell viability as well as for elongating cells
and stomata. The commonality of traits in halophytes and glycophytes is
emphasised by Flowers et al. (2010), particularly the ability for osmotic
adjustment.
Osmotic adjustment is of course essential for adapting plants to soils of
low water potential, but may bring penalties in terms of carbon allocation for
the rapidly growing phase of rapidly growing plants such as wheat and barley
(Munns, 1988). Compatible solute synthesis comes with an energy cost and
hence involves a potential growth penalty (Munns, 2005; Munns and Tester,
2008). For example, in leaf cells, approximately 7 moles of ATP are needed to
accumulate 1 moles of NaCl as an osmoticum, whereas the amount of ATP
required to synthesise 1 mole of an organic compatible solute is an order of
magnitude higher (Raven, 1985). The synthesis of these compounds occurs at
the expense of plant growth, but may allow the plant to survive and recover
from the period without water.
E. DIFFERENCES—SALT SPECIFIC
The critical issue for tolerance of a saline soil is to keep Naþ and Cl
concentrations in the cytoplasm below toxic levels. Naþ must be partitioned
within cells so that concentrations in the cytoplasm are kept low, possibly as
low as 10–30 mM (Munns and Tester, 2008). However, the concentration at
which Naþ becomes toxic is not well defined. In vitro studies showed Naþ
10 RANA MUNNS
starts to inhibit most enzymes at concentrations approaching 100 mM,

although some enzymes are sensitive to lower concentrations (reviewed in
Munns and Tester, 2008).
The concentration at which Cl becomes toxic in the cytoplasm is even less
well defined, but is probably similar to that for Naþ (Munns and Tester,
2008; Teakle and Tyerman, 2010).
Mechanisms for keeping cytoplasmic concentrations of Naþ and Cl
below toxic levels are of two main types: those minimising the entry of salt
into the root and its transport through the plant, and those minimising the
concentration of salt in the cytoplasm by sequestration in vacuoles.
Roots must exclude most of the Naþ and Cl dissolved in the soil solution
or the salt will gradually build up with time in the shoot to toxic levels. Plants
transpire about 50 times more water than they retain in their leaves (Munns,
2005). If a plant lets in only 1/50 of the salt in the soil solution, that is, it
excludes 98%, the concentration of salt in the shoot as a whole would never
increase over that in the soil and the plant could survive indefinitely in saline
soil (Munns, 2005).
Most plants in fact do exclude about 98% of the salt in the soil solution,
allowing only 2% to be transported in the xylem to the shoots. A comparison
between cereal genotypes with contrasting rates of Naþ uptake when grown
in 50 mM NaCl showed that bread wheat excludes more than 98% of the Naþ
in the soil solution, and the concentrations do not build up in leaves to more
than 50 mM (summarised in Munns, 2005). The concentrations of Naþ in the
xylem were 1–2 mM. Barley, however, had higher concentrations in the
xylem, about 3 mM, which means it excludes less than 98% of the Naþ in
the soil solution, and concentrations in leaves reach very high levels, up to
500 mM (e.g. Rawson et al., 1988). Cl exclusion also differs between species,
being 1–2 mM for bread wheat and about 5 mM for barley (Munns, 2005).
This high degree of exclusion of Naþ and Cl from the leaves is achieved
by (i) tightly controlled uptake from the soil, the epidermis of the roots
forming a virtual ‘barrier’ to entry of salt into the roots (Läuchli et al.,
2008) and (ii) regulated movement in the xylem, by controlled loading of
Cl into the xylem (Tregeagle et al., 2010) or by retrieval of Naþ as it moves
in the transpiration stream to the leaves (James et al., 2006).
Export from leaves in the phloem could conceivably help to maintain low
salt concentrations. However, there appears to be relatively little retransloca-
tion of salt from leaves, in relation to the import in the transpiration stream
(Munns, 2005). For example, estimates of xylem and phloem fluxes indicated
that in barley, phloem export from a leaf was only about 10% of the import in
the xylem (Munns et al., 1986). Exclusion of salt from the phloem ensures
that salt is not redirected to growing tissues of the shoot. Although salt that is
loaded into the phloem in lower leaves may be translocated down to the
roots, the phloem from younger leaves may go up to the meristematic and
elongating tissues in the shoot. As shown by C14 urea feeding studies, lower
leaves feed carbon to the root, upper leaves feed the shoot apex and mid-
position leaves feed both shoot apex and root (Layzell et al., 1981).
Low rates of ion accumulation in leaves could also be influenced by the
shoot:root ratio, or the relative growth rates, either of which would affect the
rate at which the ion accumulates in shoots (Munns et al., 2006).
Naþ exclusion from leaves is associated with salt tolerance in cereal crops
including rice (Asch et al., 2000; Ul Haq et al., 2010), durum wheat (Munns
and James 2003), bread wheat (Cuin et al., 2009, 2010), barley (Shavrukov
et al., 2010), pearl millet (Krishnamurthy et al., 2007) and wild relatives such
as Hordeum species (Garthwaite et al., 2005), tall wheatgrass (Colmer et al.,
2006) and Triticum tauschii (Schachtman et al., 1991).
Most studies focus on Naþ rather than Cl, as genotype comparisons
within most species link Naþ accumulation in leaves negatively with salt
tolerance. This is particularly so for species that accumulate high concentra-
tions of Cl and not Naþ in leaves, such as soybean, woody perennials such
as avocado, and species that are routinely grown on Cl-excluding root-
stocks such as grapevines and citrus. For these species, Cl toxicity is more
likely to occur than Naþ toxicity. However, this statement does not imply
that Cl is more metabolically toxic than Naþ, rather these species are better
at excluding Naþ from the leaf blades than Cl. For example, Naþ does not
increase in the leaf blade of grapevines until after several years of exposure to
saline soil, then the exclusion within the root, stem and petiole breaks down,
and Naþ starts to accumulate in the leaf blade, whereas leaf blade Cl
concentrations increase progressively (Prior et al., 2007). Cl exclusion is
critical for production of certain species on saline soil, and Cl excluding
rootstocks are used in the citrus and grapevine industry (Storey and Walker,
1999; Teakle and Tyerman, 2010; Tregeagle et al., 2010).
The mechanisms of Naþ transport at the molecular, electrophysiological
and whole plant level have been summarised by Tester and Davenport (2003)
and Munns and Tester (2008). In contrast to Naþ, mechanisms of Cl
transport in plants are poorly understood (Teakle and Tyerman, 2010).
IV. ADAPTATIONS, TRAITS AND QTLS
With the knowledge of adaptations and the mechanisms behind them, quan-
titative traits can be identified or hypothesised. These traits can be physio-
logically complex but easily measured, like leaf shape or time of flowering,
12 RANA MUNNS
which can be measured with a ruler or a visual record. They can be complex
but need expensive equipment like water use efficiency which is measured as
carbon isotope discrimination by a mass spectrometer. Or they could be
physiologically simple but difficult to measure like leaf rolling.
If a trait can be easily and inexpensively measured as a quantitative
measurement (not like presence or absence of leaf rolling), then a large
number of genotypes can be screened. If there is genetic variation, then a
QTL analysis can be performed, by crossing the contrasting parents, screen-
ing the F2 generation for the trait (the phenotype) and mapping with molec-
ular markers. The QTL should be validated in a different population. If the
QTL is robust and repeatable in different genetic backgrounds, fine mapping
can be attempted with the aid of a model species, and a candidate gene may
be revealed. If not, the original linked marker can be used for breeding
purposes.
A. DROUGHT
When considering traits that are useful in plant breeding in drought-prone

environments, the term ‘drought tolerance’ is not appropriate. It is not useful
that a plant tolerates a drought, that is, it does not die; it is only useful that it
continues to develop and produce grain. Desiccation tolerance is only appli-
cable to plants that can undergo and survive a prolonged dry season, many
months or many years.
‘Drought resistance’ is a nebulous term. There are no units for ‘drought
resistance’. Likewise, there are no units for ‘drought tolerance’. It is more
effective to ask, given a fixed and limiting water supply, what is the best
growth rate or maximum biomass that can be produced. This focuses on the
efficiency of water use, that is, water use when the plants have water, not
when they do not (Passioura and Angus, 2010).
Traits for grain production under drought have been comprehensively
reviewed and discussed by Richards et al. (2002, 2010), Reynolds et al.
(2007) and Reynolds and Tuberosa (2008). Richards et al. (2010) reviews
the key ‘best-bet’ traits for water-limited environments. These are sum-
marised in Table I.
Measurable adaptive characters can be considered in four classes: whole
plant growth and development, root architecture, leaf architecture and lon-
gevity and biochemical properties. Some of these traits have QTLs that are
reliable and useful for breeding, and some have candidate genes; however,
most have only a phenotype (see Table I).
Traits involving whole plant growth and shoot development are thorough-
ly reviewed by Richards et al. (2010). These are:
TABLE I
Summary of the Most Important Traits and the Selection Method for Improving Yield
of Temperate Cereals in Water-Limited Environments
Markers or
Environment in genomic
which trait selection regions Most efficient
Trait is most effective identified selection method
Seedling Favourable Yes Phenotype
establishment and marker
Shoot vigour Favourable Yes Phenotype
Root vigour Favourable Yes Phenotype
Root depth Favourable soil No Phenotype
moisture at
depth
Transpiration Favourable Yes Phenotype
efficiency (CID)
Stem carbohydrate Favourable Yes Phenotype
remobilisation
(WSC)
Tillering Favourable Yes Phenotype
or marker
Glaucousness Favourable Yes Phenotype
Leaf rolling Favourable No Phenotype
Floret sterility Non-droughted No Phenotype
Reproduced with permission from Richards et al. (2010); doi 10.1071/FP09219.
. seedling establishment
. shoot vigour
. tiller regulation
. phenology and early flowering
. floret sterility
Traits involving root vigour and architecture have attracted less interest,
largely because they are so difficult to measure. Root system vigour describes
the variation in rate of root growth that results in the capture of greater
volumes of soil water and nutrients (Palta and Watt, 2009). Deeper roots can
access more water, maintain high stomatal conductance and hence photo-
synthesis, and are indicated by cooler canopies (Lopes and Reynolds, 2010).
Root systems research has been hampered by the difficulty and time-consum-
ing methods of measurement, and so molecular markers would provide a new
impetus for their selection and use in breeding. The fast-growing grass
Brachypodium distachyon has a small and fully sequenced genome and as a
model for wheat and other cereal crops should provide a breakthrough in the
use of genomics and other genetic technologies (Watt et al., 2009)
14 RANA MUNNS
Traits involving leaf architecture have been reviewed by Richards et al.

(2010). The main traits are:
. maintenance of green leaf area (also called stay green)

. osmotic adjustment
. leaf rolling
. glaucousness (leaf wax properties to reflect light)
Biochemically based properties, which are complex but for which there are
markers (Richards et al., 2010) or candidate genes, are:
. transpiration efficiency
. stem carbohydrate storage and remoblisation
. osmotic adjustment and compatible solutes
B. SALINITY
Traits that increase productivity on dry land would also increase productivity
in saline land. Salinity as well as drought is a common feature of arid and
semi-arid zones, so early vigour, rapid phenology, avoidance of floret sterility
and transpiration efficiency would be as important in saline soils as well as in
dry soils.
In saline as in dry land, the term ‘salt-tolerance’ is not a useful concept for
crop growth and yield. A more useful concept is the efficiency of use of the
available water. The term may be applicable only for perennial species. In
most evergreen shrubs, survival and not growth performance is the primary
outcome of ‘salt-tolerance’ mechanisms (Tattini et al., 2009).
C. SALT-SPECIFIC TRAITS
Traits for saline soils were summarised by Colmer et al. (2005) and listed in
Table II. The first traits listed are salt specific, the last four in common with
drought.
For Naþ exclusion, and maintenance of high Kþ/Naþ ratios in leaves,
HKT genes are considered important in the regulation of Naþ and Kþ in
higher plants, and in mediating salinity tolerance in plants (Horie et al., 2009;
Munns and Tester, 2008; Pardo, 2010). HKT genes are important for cellular
Naþ and Kþ homeostasis and some family members are responsible for
TABLE II
Key Traits for Salt Tolerance in Wheat and Barley. Recommendations on Approaches, Plant Stages and other Considerations for Screening
Amenable to
large scale
Trait Measurement Plant stage Comment evaluation?
þ þ
1 Na ‘exclusion’ Leaf blade Na At defined leaf age (days The most useful trait; Yes
concentration after emergence) and requires only small
stage of plant tissue samples (i.e.
development (leaf non-destructive); Cl
number) ‘exclusion’ is much less
studied, and for wheat
showed little
correlation with
genotypic differences
in salt tolerance
2 Kþ/Naþ discrimination Leaf blade Naþ and Kþ As above Leaf Naþ alone is Yes
sufficient
3 Sheath retention of ions Naþ and/or Cl As above Not widely studied Yes
concentration in
sheath and blade
4 Tissue tolerance Leaf injury, coupled with Seedling stage Laborious to phenotype No
Naþ and/or Cl
accumulation
5 Ion partitioning into Leaf blade Naþ Pre- and post-stem See also (1) above Yes
different-aged leaves concentration of elongation
different leaves
6 Osmotic adjustment Turgor or concentration Seedling stage Assessing tissue water Turgor—no
of specific solutes relations is laborious; Solutes— yes
screening organic
solutes requires a high-
throughput analytical
system
(continues)
Table II (continued )
Amenable to
large scale
Trait Measurement Plant stage Comment evaluation?
7 Enhanced vigour Area of first leaves Seedling stage Enhances water uptake Yes
early in season, and
reduces evaporation
from soil
12
8 Water use efficiency C:13C discrimination Late vegetative stage Reduces water uptake Yes
(low or 13C) later in the season;
might be useful for
transient salinity only
9 Early flowering Flowering date Flowering May reduce yield Yes
potential
Reproduced with permission Colmer et al. (2005); doi 10.1071/EA04162.
Naþ exclusion from leaves. The transporters encoded by these HKT genes
can remove Naþ from the xylem and result in lower Naþ in leaves and a
higher K/Naþ ratio which is important to maintain ion homeostasis of the
cytoplasm (Munns and Tester, 2008). The Arabidopsis gene AtHKT1;1 is a
Naþ transporter that limits Naþ transport from root to shoots by removing
Naþ from the xylem (Davenport et al., 2007). Overexpression of this gene
with a stele-specific promoter reduced the rate of transport of Naþ to the
leaves and improved the salt tolerance of Arabidopsis (Moller et al., 2009).
Although dicots have only one family member, monocots have several, and
there are nine in rice, wheat and barley (Huang et al., 2008). HKT members
of sub-family 1 have a much higher selectivity for Naþ over Kþ than do
members of sub-family 2 (Pardo, 2010; Hauser and Horie, 2010).
Major genes with QTLs have been found for five Naþ excluding genes—
Nax1 and Nax2 in durum wheat, Nax3 and Nax4 in barley and Kna1 in bread
wheat as described below.
In durum wheat, Nax1 and Nax2 enhanced removal of Naþ from the
xylem, leading to low Naþ concentrations in leaves (James et al., 2006).
Nax1 removes Naþ from the xylem in roots and the lower parts of leaves,
the leaf sheaths, while Nax2 removes Naþ from the xylem only in the
roots (James et al., 2006). Nax1 has a unique phenotype of a high sheath:
blade ratio of Naþ concentration. Nax2 has the same phenotype as Kna1,
the QTL for Naþ exclusion and enhanced K/Naþ selectivity in bread
wheat, Triticum aestivum (Dvořák et al., 2004). Nax2 was shown to be
homoeologous to Kna1 in T. aestivum, namely TaHKT8 (TaHKT1;5) (Byrt
et al., 2007).
Modern dwarf rice is less tolerant of salinity than wheat, but the original
landraces Pokkali and NonaBokra are relatively tolerant. This is due largely
to the presence of an orthologue of TmHKT1;5-A, namely OsHKT1;5 (Ren
et al., 2005).
However, no QTL for Naþ exclusion has been found on chromosome 4,
the syntenic region to wheat chromosome 2A containing the Nax1 gene. Yet
the gene sequence for HKT1;4 is present on the rice genome (Huang et al.,
2006). It is possible that this gene is silenced in rice, or for whatever reason
its expression is prevented, leading to the interesting idea that transformation
of rice with TmHKT1;4 might increase its salt tolerance. This might be
particularly beneficial when the soil is flooded and insufficient oxygen
reaches the roots, despite the aerenchyma, to maintain ion transport activity
in roots.
The family of HKT genes are not the only genes controlling Naþ transport
to the shoot. Studies with Na22 showed that durum wheat lacking the Nax
18 RANA MUNNS
genes was able to withdraw about 50% of the Na22 flowing to the shoot (James
et al., 2006). The presence of either Nax1 or Nax2 increased this withdrawal
from the xylem to about 90%. QTLs for Naþ exclusion in leaves of barley
revealed two other major genes, named Nax3 (Shavrukov et al., 2010) and
Nax4 (Rivandi et al., 2011). The latter locus contains a SOS3 homologue.
Efflux of Naþ across the plasma membrane to the apoplast and ultimately to
the soil solution is by Naþ/Hþ antiporters (Blumwald et al., 2000; Pardo, 2010;
Pardo et al., 2006). SOS1 exchanger is regulated through the SOS2–SOS3
kinase complex in Arabidopsis, and a similar pathway is important in salt
tolerance of this (Shi et al., 2003) and other species (reviewed by Pardo, 2010).
NHX1, the vacuolar Naþ/Hþ antiporter controls Naþ sequestration in
cells (Blumwald et al., 2000), leading to the concept of ‘tissue tolerance’—the
ability of some species to tolerate high concentrations of Naþ in their leaves,
presumably because it is largely sequestered in the vacuole (Flowers and Yeo,
1986). Undoubtedly, there are genotypic differences, but it is difficult to
measure. We found that there was a relation between the Naþ concentration
at which the leaf died, or lost 50% of its chlorophyll content, but that this
quantitative relationship differed for leaves of different ages (R. Munns and
R. James, unpublished data). This means that just linking the increase in Naþ
concentration with a decrease in chlorophyll is inconclusive.
In the list in Table II, Cl exclusion as a trait is not mentioned, because the
chapter was focused only on wheat and barley, for which species there is no
evidence of Cl toxicity, and no correlation between genotypic differences in
Cl uptake and salinity tolerance. Genes important in the regulation of Cl
transport in saline soil are reviewed by Teakle and Tyerman (2010). No
QTLs for these genes have been identified.
A decrease in leaf chlorophyll content is an indicator of tolerance, and an
easily measurable phenotype, but there is a problem of assigning cause. A
decrease in chlorophyll could be due to lower tolerance of tissue Naþ or Cl
concentrations, or premature senescence caused by signals from the roots.
Maintenance of a high Kþ concentration in leaves and roots, and a ratio of
K /Naþ greater than 1, is also crucial for salt tolerance (Hauser and Horie,
þ
2010; Shabala and Cuin, 2008. The ability of a plant to retain Kþ in the roots
exposed to high NaCl concentrations is a critical feature of tolerance in many
species (reviewed by Shabala and Cuin, 2008), and genetic variation in NaCl-
induced Kþ efflux from roots correlates with genetic variation in grain yield
of barley (reviewed by Shabala and Cuin, 2008) but not of wheat (Cuin et al.,
2009). In wheat, measurements of shoot sap Kþ concentration in non-sali-
nised plants, along with the changes of chlorophyll concentration in a given
leaf after 6 weeks of NaCl treatment correlated with % grain yield in saline
soil and were proposed as an efficient screening tool (Cuin et al., 2010).
D. TRAITS IN COMMON WITH DROUGHT
The last four traits listed in Table II are in common with drought.
Osmotic adjustment occurs readily because of the high uptake of Naþ and

Cl , and in species like wheat and barley is regulated so that it does not
become toxic. Turgor is perfectly maintained (Boyer et al., 2008). Compati-
ble solutes are needed to balance the Naþ and Cl in the vacuole as their
concentration there may be only 30 mM, as mentioned earlier in this chapter
(Section III.E).
If Naþ and Cl are sequestered in the vacuole of a cell, organic solutes that
are compatible with metabolic activity even at high concentrations (hence
‘compatible solutes’) must accumulate in the cytosol and organelles to bal-
ance the osmotic pressure of the ions in the vacuole (Flowers et al., 1977).
The compounds that accumulate most commonly are sucrose, proline, and
glycine betaine, although other molecules can accumulate to high concentra-
tions in certain species.
In many halophytes, proline or glycine betaine occurs at sufficiently high
concentrations in leaves (over 40 mM on a tissue water basis) to contribute to
the osmotic pressure (over 0.1 MPa) in the cell as a whole (Flowers et al.,
1977). In glycophytes, the concentrations of compatible solutes that accumu-
late are not so high, on the order of 10 mM, but if partitioned exclusively to
the cytoplasm, they could generate a significant osmotic pressure and func-
tion as an osmolyte. At low concentrations, these solutes presumably have
another role, perhaps in stabilising the tertiary structure of proteins, and
function as osmoprotectants (Rhodes et al., 2002).
Most of the other traits in Table II are covered in the above section on
‘Drought’ (Section IV.A). Not listed in that table are root properties and
floral sterility.
1. Root properties
Durum wheat (Triticum turgidum L. ssp. durum Desf.) is relatively salt
sensitive compared to bread wheat (T. aestivum L.), and yields poorly on
saline soil (Munns and James, 2003). Field studies indicate that roots of
durum wheat do not proliferate as extensively as bread wheat in saline soil.
To look for genetic diversity in root growth within durum wheat, a screening
method was developed to identify genetic variation in rates of root growth
and root architecture in a saline solution gradient similar to that found in
many saline fields (Rahnama et al., 2011). Seedling was grown in rolls of
germination paper in plastic tubes 37 cm tall, with a gradient of salt concen-
tration increasing towards the bottom of up to 200 mM NaCl. A NaCl
concentration of 150 mM at the bottom was found suitable to identify
20 RANA MUNNS
genetic variation in seminal axile root length as well as branch root length.
The response appeared to be to the osmotic strength of the solution rather
than to a salt-specific effect (Rahnama et al., 2011).
2. Sterility
Ghanem et al. (2009) showed that the failure of the tomato inflorescence to
develop normally under salt stress was best explained in terms of altered
source–sink relationships than accumulation of ions to toxic concentrations.
Using laser ablation inductively coupled plasma mass spectrometry micro-
analysis to measure Naþ concentrations is reproductive tissues including the
tapetum, they concluded that the decrease in soluble carbohydrate was the
most likely cause of the sterility.
E. QTL ANALYSIS AND GENE DISCOVERY
The significance of knowing the essential traits is to use this information for
gene discovery, and one way is via a QTL. For this, an unambiguous and
quantitative phenotype, and an experimental design that is realistic and
reproducible are needed.
Genetic technologies for identifying genes important in environmental
stress responses have been summarised by Papdi et al. (2009). These are as
follows: traditional mutagenesis (using forward or reverse screens), insertion
mutagenesis (forward or reverse screens), QTL mapping and cDNA trans-
formation (forward screen) (summarised in Figure 7 of Papdi et al., 2009).
All require a physiologically based genetic analysis at some stage (Papdi
et al., 2009). QTL mapping requires a quantitative physiologically based
screen, although just biomass production can be successful.
All studies require a sound experimental design that can quantify growth
responses under water deficit or salinity stress, not easy in the laboratory or
when using genetic material that has different intrinsic growth rates or
phenology.
V. GROWTH STUDIES AND EXPERIMENTAL DESIGN
A. WATER STRESS
The search for the best medium for growing plants to impose a controlled
water deficit has been going on for decades, without a clear resolution. There
is no perfect medium—all have limitations including pots containing soil
(Munns et al., 2010).
Soil drying is hard to control especially when comparing genotypes of

different vigour or rates of development. Even a drying soil, as well as being
very difficult to maintain at a uniform and constant water potential through
the whole soil profile, may exert specific effects; for example, transmission of
nutrients through the soil will be reduced at low soil water potentials.
Another problem with soil is drainage, and easily become saturated at the
bottom (Passioura, 2006). Soils should drain quickly; otherwise, the ‘con-
trols’ are waterlogged. The addition of perlite or vermiculite, or the use of
‘inorganic soils’ such as fritted or calcined clay can overcome the problems of
soil with a high clay content, which does not drain quickly or a predomi-
nantly sandy soil which holds little water and releases it quickly as the soil
dries.
Hydroponics avoids problems of drainage. A variety of non-ionic osmo-
tica have been used to mimic a decrease in soil water potential, such as
mannitol. However, a percentage of these small molecules enter roots and
move in the xylem to the shoots, either through cracks in roots or through
membranes that are not completely impermeable to neutral solutes of this
size. High-molecular-weight PEG (MW 6000) has been examined in many
early studies that attempted to impose a controlled water deficit. Its main
problem is its viscosity that decreases O2 movement to roots so that the roots
become O2 deficient. The latter can be overcome by bubbling with O2 rather
than air; however, the experiments must be limited to a short period of time
as the PEG can enter the roots and reduce the hydraulic conductivity (Munns
et al., 2010). Concentrated mixed salts, such as the macronutrients used in
Hoagland’s solution, are preferable to NaCl as plants are less likely to take
up any one ion to toxic concentrations (Munns, 2010a, in PrometheusWiki).
When hydroponics are used to simulate a ‘drought’ or soil water deficit,
the effect on plant growth may be quite different, and a lot less, than with a
drying soil. Hydroponics ensure that there is no nutrient deficit, but in a dry
soil the access to nutrients decreases and the plants may suffer N or P
deficiency (summarised in Munns et al., 2010).
Experiments in drying soil are difficult when comparing genotypes of
different sizes such as mutants or transformants whose size or relative growth
rate is less than that of the wild type, or when comparing modern versus old
cultivars whose size has been reduced by the introduction of dwarfing genes.
When genetically modified lines are compared with their wild type, the soil
water status must be measured and their performance compared at the same
soil water potential; otherwise, the small plants use the soil water more slowly
and stay greener longer, and appear to be more ‘drought tolerant’ when
photographs are taken. Rewatering should be avoided, as it favours the small
plants.
22 RANA MUNNS
B. SALINITY
Three treatment approaches can be taken (Munns, 2010b, in Prometheus-

Wiki). The most common and convenient one is solution culture, which can
be supported by solid material such as fine gravel or high-density plastic
beads. A recirculating nutrient solution using a modification of the original
Hoagland’s solution (Munns and James, 2003) is applied using aeration in
pots or subirrigation in tanks. A second method is sand culture, when the
sand is irrigated with Hoagland’s nutrient solution. A third method is to use
soil as the medium, which is likely to best mimic field conditions, but in small
pots the soil needs to flushed periodically with salt-nutrient solution (making
is similar to sand culture but with reduced drainage) or rewatered by repla-
cing the water transpired but in this case it creates pockets of low-salinity soil
in high-salinity background (Munns, 2010b).
Experiments should be conducted over a reasonable time period, and the
salt increased in gradual steps to avoid severe osmotic shock. Plant responses
in the short term are primarily osmotic, and only in the longer term (days,
weeks or months, depending on the species) does the salt rise to toxic
concentrations in leaves and the salt-specific effect is seen (Munns, 2002).
Experiments that have compared hydroponics and soil at a range of
salinities have found quite different qualitative and quantitative differences.
Differences between two barley cultivars in growth, moisture content and
Naþ accumulation were not apparent in hydroponics but significant differ-
ences occurred in soil (Tavakkoli et al., 2010) due to salt moving in the soil as
it was watered, probably.
Ca2þ deficiency is an artefact generated in hydroponics. Kopittke et al.
(2010) provide a comprehensive analysis of the effect of any one cation on the
activities of other cations in solution and the effect on plant growth rates.
Short-term (2d) growth of cowpea seedling to Cl salts of Naþ, Kþ, Ca2þ
and Mg showed that growth was poorly related to activities in the bulk
solution but closely related to activities at the outer surface of the plasma
membrane. The addition of Mg Naþ or Kþ resulted in Ca2þ deficiency in
roots at Ca2þ concentrations less than 1.6 mM.
C. EXPERIMENTS TO DISTINGUISH WATER STRESS FROM SALT-SPECIFIC

EFFECTS
To distinguish the effects of water stress, as in a dry soil, from salt-specific

effects, can most easily be done in a time course. The osmotic effect of the soil
salinity, as does the dehydrating effect of the soil, has an immediate effect on
growth rate (Fig. 2). The effect of toxicity on the older leaves takes time to
show up (Munns, 2002; Munns et al., 1995). This concept led to Rajendran
et al., 2009 showing that it could be quantified within 5–7 days by non-
destructive leaf area measurements, and genotypic differences in Triticum
monococcum were distinguished.
The ‘salt transcriptome’ was studied over time in a salt-tolerant Poplar
species along with physiological changes, in order to identify genes important
in the acclimation to stress—rather than just the response to the osmotic
shock (Brinker et al., 2010). Over 24 h, three distinct phases of salt stress:
dehydration (due to water withdrawal and osmotic shock), salt accumulation
(rapid uptake) and osmotic restoration (regaining osmotic or ionic homeo-
stasis). The initial dehydration was more marked in leaves than in roots and
resulted in changes in transcript levels of mitochondrial and photosynthetic
genes indicating adjustments of energy metabolism. Stress-specific genes
responded only when leaves had osmotically adjusted by salt accumulation.
Another way of distinguishing osmotic from salt-specific effects is to
compare the growth of plants in NaCl with that in other salts, with mixed
salts, or in non-ionic media.
A variety of non-ionic osmotica have previously been used to mimic a
decrease in soil water potential, including mannitol, melibiose and sorbitol.
However, all these small molecules eventually enter roots and move in the
xylem to the shoots because membranes are permeable to neutral solutes of
this size (reviewed in Munns et al., 2010). High-molecular-weight PEG has
been examined in many early studies that attempted to impose a controlled
water deficit. Its main problem is that its viscosity limits O2 diffusion so that
the roots become O2 deficient (Verslues et al., 1998). When PEG solutions
were bubbled with O2 (not vigorously with air, which breaks roots), the rate
of root growth increased and the partial pressure of O2 at the root tip
increased (Verslues et al., 1998). A problem with solution culture is that
lateral roots break when the solution is changed and the roots are unsup-
ported, and solution enters through the damaged junction between a lateral
root and the seminal axis. This allows a pulse of salt into the plant (own
observations) and also provides a mode of entry for PEG or other osmotica
(Munns et al., 2010). This problem can be overcome by using solid media
such as quartz gravel, which supports the roots when the solution is removed
or aerated (e.g. Munns and James, 2003).
Mixed salts, for example, high concentrations of the macronutrients in
Hoagland’s solution, are a better osmotica than any of the above, as their
rate of uptake is tightly regulated by transporters that have evolved to deal
with variable concentrations in soils, and they do not support bacterial
growth. Concentrated macronutrients were shown by Termaat and Munns
(1986), Rengasamy (2010a) and Tavakkoli et al. (2010) to be a good
24 RANA MUNNS
surrogate for water deficit caused by osmotic stress. Figure 2 shows that
growth of wheat was reduced by 7 dS/m of NaCl, about 0.25 MPa, more
than by mixed salts (derived from concentrating the macronutrients of Hoag-
land’s solution) of the same total concentration. However, at the higher
concentration of 30 dS/m or 1.1 MPa, there was less difference, indicating
the osmotic effect became the overriding effect on growth at the higher
osmotic strength. The figure also illustrates that differences did not show up
for 25 days of treatment. In other more salt-sensitive species, which do not
have such a tight regulation of Naþ or Cl transport to the leaves, differences
will show up earlier.
To distinguish Naþ from Cl toxicity is surprisingly difficult. One might
consider that comparing growth rates in different salts should be informative.
For example, the specific effect of Cl could be tested by comparing NaNO3 or
Na2SO4 versus NaCl, and specific effect of Naþ by comparing CaCl2 or KCl
versus NaCl. These experiments have been tried for over 50 years and have not
produced unambiguous results. The main reason is that the osmotic effect
overwhelms the specific salt effect (Kopittke et al., 2010; Rengasamy, 2010a).
Furthermore, the effect of any single salt can be toxic, more so than NaCl.
Plants have evolved to exclude Naþ and Cl, as they are the most prolific
soluble salts in the soil, and when presented with high concentrations of NO3,
SO4 or Kþ, they take up large amounts. Ca2þ salts are least toxic (Rengasamy,
2010a) but they do not dissociate as fully in solution as other salts and allowance
must be made for their lower osmotic strength for a given concentration.
VI. PHENOTYPING
Many of the traits for drought and salt tolerance have a clear phenotype, and
many of these have molecular markers, if not perfect markers derived from
the candidate genes.
However, some do not or the technique is too expensive and slow, for
example, transpiration efficiency. In this case, a measurement of plant
growth rate is more cost effective. Also, we may have insufficient knowledge
of traits and their interactions, and plant breeders still select on yield, so a
measurement of growth rate, biomass or yield, empirical though it is, may be
the most successful.
Selection techniques thus can make use of modern ‘phenomics’ tools. Non-
destructive, high-throughput crop evaluation in controlled environments and
the field is being developed (Berger et al., 2010; Furbank, 2009). This enables
a correlation of gene function with plant performance and environmental
response, with high resolution and speed.
Infrared thermography as a surrogate for measurements of transpiration

and photosynthetic response to stress shows great promise in high-through-
put plant phenomics and can be used both in controlled environments
(Sirault et al., 2009) and in the field (Jones et al., 2009). The power of this
approach is that it allows high-throughput screening at the seedling level to
be validated at the canopy level in a field environment, using the same
genotypes and tools.
Spectroscopic techniques that quantify photosynthetic performance and
can be used at the leaf and canopy level are described briefly by Furbank
(2009) and Munns et al. (2010). Spectral reflectance and absorption measure-
ments can potentially provide plant phenomics with a tool to non-invasively
delve into plant function and chemical composition scalable from the tissue
to the canopy level. For example, high-throughput chlorophyll fluorescence
imaging using a 9 8 grid array can rapidly identify mutants in photorespi-
ration (Badger et al., 2009).
Determining root function in soil and screening for optimising root struc-
ture and growth has long been a challenging field (reviewed by Gregory et al.,
2009). Access to water at depth for cereal species growing on stored soil
moisture is of great importance for drought tolerance and screening of
model species to elucidate genes responsible for root characters is underway.
Recently developed small, short-lifecycle crop models more appropriate to
cereal species are excellent systems for phenomic screening (Watt et al., 2009).
Deeper roots can access more water, maintain high stomatal conductance
and hence photosynthesis and be indicated by cooler canopies (Lopes and
Reynolds 2010), a phenotype less laborious than soil coring (Watt et al.,
2005). Infrared thermography (Jones et al., 2009), canopy greenness or
carbon isotype discrimination have the potential to provide a non-destructive
phenotype (Richards et al., 2010). Looking ahead, new non-destructive
methods to measure soil moisture should become available such as ground
penetrating radar and electrical resistance thermography, and become suffi-
ciently sensitive to detect genotypic differences.
Another application of modern phenomics is in gene discovery. A signifi-
cant proportion of even the Arabidopsis genome is annotated as ‘gene of
unknown function’ or annotated using only loose sequence homology-based
clues.
Reverse genetic approaches to disrupt gene function often result in the
unsatisfactory conclusion ‘no visible phenotype’ (Furbank, 2009). The recent
developments of chlorophyll fluorescence techniques combined with infrared
thermography provide information on photosynthetic and stomatal func-
tions that can reveal new knowledge on the functions of key genes that
ultimately control plant growth.
26 RANA MUNNS
VII. CONCLUSIONS
The future of gene-based approaches to increasing growth and yield under

drought and salinity depends on physiological understanding and realistic
experiments.
The significance of knowing the essential traits is to use this information
for gene discovery, via a QTL. For this, an unambiguous and quantitative
phenotype, and an experimental design that is realistic and reproducible are
needed. QTLs need to be validated in different populations. Other
approaches of traditional or insertion mutagenesis (using forward or reverse
screens) and cDNA transformation (forward screen) also require good ex-
perimental design, and the information is enhanced if a trait is hypothesised.
Traits in controlled environment chambers need to be validated in the
field. The field brings many other factors—atmospheric particularly vapour
pressure deficit and temperature, and the soil will undoubtedly contain other
constraints such as poor structure physical such as compaction or chemical
such as acidity or sodicity. There are many different categories of saline soil,
arising from different mineral compositions and different types of salinisa-
tion process, all affecting plant responses to the increased osmotic pressure or
specific ion concentrations (Rengasamy, 2010b).
Salinity can be transient (Rengasamy, 2006) as can waterlogging, and the
salinity–waterlogging interaction can severely inhibit plant growth (Colmer
et al., 2005).
Drought and salinity limit the production of the world’s staple food crops.
Conventional plant breeding based on yield in target environments has
increased production; however, physiologically based approaches utilising
molecular tools to identify key genes or provide molecular markers have the
potential to take it further. Accurate and selective phenotyping will enable
the best use of mechanistic and molecular understanding of plant responses
to drought and salinity, and mechanisms of adaptation.
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Plant Adaptations to Salt and Water Stress: Differences and

CSIRO Plant Industry, Canberra, and School of Plant Biology,

Advances in Botanical Research, Vol. 57 0065-2296/11 $35.00

ha of land farmed by dryland agriculture are affected by secondary salinity to

II. WHOLE PLANT RESPONSES TO WATER AND SALT

20 7.0 dS/m NaCl 20 30.0 dS/m NaCl

III. MECHANISMS OF DROUGHT AND

The understanding of plant responses to water deficits from the physiolog-

B. LONG-DISTANCE SIGNALS CONTROLLING LEAF EXPANSION AND

Whether water status, hormonal regulation or supply of photosynthate

C. CHANGES IN LEAF MORPHOLOGY AND ROOT SYSTEM ARCHITECTURE

Hormonal control of cell division and differentiation is clear from the

Recently, the concept of phenotypic plasticity has re-emerged, partly

D. TURGOR MAINTENANCE AND OSMOTIC ADJUSTMENT

starts to inhibit most enzymes at concentrations approaching 100 mM,

IV. ADAPTATIONS, TRAITS AND QTLS

When considering traits that are useful in plant breeding in drought-prone

Traits involving leaf architecture have been reviewed by Richards et al.

. maintenance of green leaf area (also called stay green)

D. TRAITS IN COMMON WITH DROUGHT

E. QTL ANALYSIS AND GENE DISCOVERY

V. GROWTH STUDIES AND EXPERIMENTAL DESIGN

Soil drying is hard to control especially when comparing genotypes of

Three treatment approaches can be taken (Munns, 2010b, in Prometheus-

C. EXPERIMENTS TO DISTINGUISH WATER STRESS FROM SALT-SPECIFIC

To distinguish the effects of water stress, as in a dry soil, from salt-specific

Infrared thermography as a surrogate for measurements of transpiration

The future of gene-based approaches to increasing growth and yield under

Munns, R. (2010a). Drought treatments. In PrometheusWiki, Version 1. http://www.

Prior, L. D., Grieve, A. M., Bevington, K. B. and Slavich, P. G. (2007). Long-term

Schlichting, C. D. (1986). The evolution of phenotypic plasticity. Annual Review of

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