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Improving agricultural water use efficiency by

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a a b c
Ejaz Ahmad Waraich , Rashid Ahmad , M. Yaseen Ashraf , Saifullah & Mahmood
d
Ahmad
a
Department of Crop Physiology, University of Agriculture, Faisalabad, Pakistan
b
Nuclear Institute of Agriculture and Biology, Faisalabad, Pakistan
c
Institute of Soil and Environmental Sciences, University of Agriculture, Faisalabad,
Pakistan
d
Department of Agriculture Extension, University of Agriculture, Faisalabad, Pakistan

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 Acta Agriculturae Scandinavica Section B
Soil and Plant Science, 2011; 61: 291
304

REVIEW ARTICLE

Improving agricultural water use efficiency by nutrient management

in crop plants

EJAZ AHMAD WARAICH1, RASHID AHMAD1, M. YASEEN ASHRAF2, SAIFULLAH3 &

MAHMOOD AHMAD4
1
Department of Crop Physiology, University of Agriculture, Faisalabad, Pakistan, 2Nuclear Institute of Agriculture and
Biology, Faisalabad, Pakistan, 3Institute of Soil and Environmental Sciences, University of Agriculture, Faisalabad, Pakistan,
4
Department of Agriculture Extension, University of Agriculture, Faisalabad, Pakistan
Downloaded by [Wageningen UR] at 00:05 15 December 2011

Abstract
Improvement of agricultural water use efficiency is of major concern with drought problems being one of the most
important factors limiting grain production worldwide. Effective management of water for crop production in water-scarce
areas requires efficient approaches. Increasing crop water use efficiency and drought tolerance by genetic improvement and
physiological regulation may be a means to achieve efficient and effective use of water. A limited water supply inhibits the
photosynthesis of plants, causes changes of chlorophyll contents and components and damage to photosynthetic apparatus.
It also inhibits photochemical activities and decreases the activities of enzymes in plants. Water stress is one of the important
factors inhibiting the growth and photosynthetic abilities of plants through disturbing the balance between the production
of reactive oxygen species and the antioxidant defence, causing accumulation of reactive oxygen species which induce
oxidative stress to proteins, membrane lipids and other cellular components. A number of approaches are being used to
enhance water use efficiency and to minimize the detrimental effect of water stress in crop plants. Proper plant nutrition is a
good strategy to enhance water use efficiency and productivity in crop plants. Plant nutrients play a very important role in
enhancing water use efficiency under limited water supply. In this paper we discuss the possible effective techniques to
improve water use efficiency and some macronutrients (nitrogen, phosphorus, potassium, calcium and magnesium),
micronutrients (zinc, boron, iron, manganese, molybdenum and chloride), and silicon (a beneficial nutrient) in detail to
show how these nutrients play their role in enhancing water use efficiency in crop plant.

Keywords: Macronutrients, mechanisms, micronutrients, water use efficiency.

Introduction The world’s population (6000 million) of today is

expected to reach about 8100 million by 2030,
Water use efficiency (WUE) can be described on
an increase of 35%. The growing population will
various scales from the leaf to the field (Sinclair
result in considerable additional demand for food.
et al., 1984). In its simplest terms, WUE refers to the
Simultaneously, water demand from non-agricultural
ratio of yield to the water used during crop growth
sectors will keep growing in both developed and
(Waraich et al., 2008) whereas the ratio of yield to
developing countries. About 40% of the land in the
transpiration is termed as transpiration efficiency
world is under arid and semi-arid climatic conditions
(TE). Water use efficiency provides a simple way of (Gamo, 1999). An FAO analysis (FAO, 2003) of 93
assessing whether yield is limited by water supply or developing countries expects an increase of agricul-
other factors. A good understanding of crop man- tural production over the period 1998
2030 by
agement effects on WUE may provide researchers 49% in rain fed and by 81% in irrigated regions.
with opportunities to identify and select appropriate Therefore, much of the additional food production is
crop management practices for improved water use expected to come from irrigated land, three-quarters
efficiency. of which is located in developing countries. The

Correspondence: Ejaz Ahmad Waraich, Department of Crop Physiology, University of Agriculture, Faisalabad, Pakistan. Tel: 92-0336-6553081:
Fax: 92-041-9200764. E-mail: [email protected]

(Received 11 March 2010; revised 1 May 2010; accepted 4 May 2010)

ISSN 0906-4710 print/ISSN 1651-1913 online # 2011 Taylor & Francis
DOI: 10.1080/09064710.2010.491954
 292 E.A. Waraich et al.

irrigated area, in developing countries, in 1998 has Water-saving irrigation

nearly doubled since 1962. The FAO estimates that
Due to increased water scarcity, the irrigated area
the irrigated area in the selected 93 developing
is unlikely to expand in the dry-land region. Supple-
countries will only grow by 23% over the 1998

mental irrigation, the combination of dry-land farm-
2030 period. However, the effective harvested irri-
ing and limited irrigation, is an ideal choice for
gated area (considering the increase in cropping
improving crop yields (Bai & Dong, 2001). Good
intensity) is expected to increase by 34%. This
irrigation scheduling requires the timing of irrigation
means that more area needs to be brought under and the amount of water applied to match actual
irrigation to produce more crops with the present field conditions. This requires information on soil-
available water resources (Ali & Talukder, 2008). In moisture conditions at the time of irrigation and,
the dry areas, water, not land, is the most limiting when using irrigation, close cooperation among
resource for improving agricultural production. farmers to be effective. Further, water-saving
Maximization of yield per unit of water (water use irrigation depends on adopting water-saving techni-
efficiency), and not yield per unit of land (land ques in the transportation and application of the
productivity), is therefore a better strategy for dry water. On-farm WUE can be improved by moving to
farming systems. Under such conditions, more a more efficient irrigation system. There are three
efficient water management techniques must be main types of irrigation systems available: border or
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adopted (Ali & Talukder, 2008).Water deficiency furrow flood irrigation, sprinkler irrigation and drip
and low availability of nutrients often limit crop irrigation, with drip irrigation being more efficient
growth and production potential in agro-ecosystems than sprinkler irrigation and sprinkler irrigation
because most crops are sensitive to water and being more efficient than border/furrow irrigation.
nutrient deficits during different critical stages. On An investigation into the irrigation of maize showed
the other hand, excessive use of water can increase that 210 mm of irrigation applied by border irriga-
production costs and pollute the environment tion under a mulch of wheat straw gave a grain yield,
(enhancing fertilizer leaching). So improvement of total evapotranspiration and WUE of 8000 kg ha 1,
water use efficiency (also termed water productivity) 390 mm and 2.2 kg m3, respectively (Zhang et al.,
in agriculture is very important. It is widely believed 2002). Increasing the amount of irrigation to
that an increase in agricultural water use efficiency is 270 mm increased the yield to 8834 kg ha 1, but
the key approach to mitigate water shortages and to WUE decreased. Straw mulching reduced the total
reduce environmental problems. evaporation by 50 mm showing that wheat straw
This paper therefore discusses the role of mineral mulching can reduce the amount of irrigation
nutrition and possible mechanisms to improve water required by maize in the North China Plain (Zhang
use efficiency. et al., 2002). Irrigation of cotton in Xinjiang
indicated that the flowering and budding stages
Crop and soil management approaches to were the most suitable times to supply limited
enhance water use efficiency irrigation water, thus significantly improving the
WUE by 57% (Hu et al., 2002). In the northern
Enhancing agricultural WUE by crop and soil part of China, Liu et al. (2003) reported that the
management approaches refers to a farming practice yield and WUE of winter wheat under sprinkler
that is able to take full advantage of the natural irrigation conditions was increased by 28 and 48%
rainfall and irrigation facilities. The core problem and 636 m3 ha1 water was saved compared with
that water-saving agricultural research has to solve is that under border irrigation conditions. Zhang and
how to raise the water utilization rate and water use Cai (2001) conducted a cotton irrigation experiment
efficiency, that is to achieve a high yield on irrigated that demonstrated that irrigation under the surface
farmland with the minimum input of water and of a plastic mulch is an effective way to protect
in rainfed agriculture to maximize rainfall use against soil evaporation and that a mild water deficit
efficiency. What we stress here is that water-saving during the budding stage could significantly
agriculture is not simply water-saving irrigation. It is enhance lint yield and improve water use efficiency.
a comprehensive exercise using every possible water- These examples show that water-saving irrigation
saving measure in whole-farm production, including can really save water and need to be used at a
the full use of natural precipitation as well as the regional scale. However, to improve the standard of
efficient management of an irrigation network and irrigation design and irrigation efficiency requires (i)
plant mineral nutrition. The different management enforcement or regulation
requiring irrigation to be
approaches by which we can improve WUE are given designed to a standard, and (ii) education
informing
below. water users of the benefits of good design to
 Agricultural water use efficiency 293

encourage them to expect a high standard of design The highest WUE of 4.4 kg m3 for biomass and
from irrigation equipment suppliers (Zhou, 2003). 2.2 kg m3 for grain yield was achieved when N
fertilizer was added and plant density was kept low.
Terracing and contour farming
Plant mineral nutrition
With frequent farming activities and a high degree of
cultivation, sloping land with an angle of 10
258 is Proper nutrition is the basic need of every living
highly susceptible to soil erosion. Cultivation on organism. There are now 17 elements which are
such slopes can lead to erosion of 0.43 cm and considered essential for plants to complete their life
48 t ha1 of fertile topsoil (Wei et al., 2000). cycle. These essential plant nutrients are divided into
Changing such sloping land into contour terraces two categories; macronutrients and micronutrients.
prevents water and soil erosion, raises land quality Macronutrients include carbon (C), hydrogen (H),
and grain yield. Sloping land with an angle of 6
108 oxygen (O), nitrogen (N), phosphorus (P), potas-
can be improved by planting crops along the contour sium (K), calcium (Ca), magnesium (Mg) and
using a 0.5-m deep and 1-m wide trench or ridge to sulphur (S). Micronutrients are zinc (Zn), copper
conserve soil and water, improve soil fertility and (Cu), iron (Fe), manganese (Mn), boron (B),
facilitate sustainable development. In the semiarid molybdenum (Mo), chlorine (Cl) and nickel (Ni).
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Loess Plateau, the building of level terraces has Although silicon (Si) is not essential, it is considered
enhanced water infiltration, raised the rainfall utili- as a beneficial plant nutrient. Adequate nutrition is
zation rate and created high-yielding farmland while essential for the integrity of plant structure and key
also conserving the soil and water. Combined with physiological processes such as light interception
other agricultural techniques, it has played a major by chlorophyll (N, Mg), energy for carbohydrate
role in increasing the productivity and sustainability build-up (P), and osmotic regulation of the stomata
of the region (Deng et al., 2000). (K), among others. Therefore, a well-nourished
plant is expected to produce more biomass per unit
of transpired water than an under-nourished one.
Mulching
Radin and Mathews (1989) also found that N and P
Mulching with crop residues can improve water use deficient plants strongly reduced the hydraulic con-
efficiency by 10
20% through reduced soil evapora- ductivity of the root cortical cells. These plant
tion and increased plant transpiration. Mulching nutrients are not only required for better plant
with crop residues during the summer fallow can growth and development, but also helpful to improve
increase soil water retention (Feng, 1999). Straw agricultural WUE. Increasing evidence suggests that
mulching can be easily implemented by local farmers mineral-nutrient status of plants plays a critical
because material is easily accessible, low cost and role in enhancing water use efficiency by proper
does not contaminate the soil. Combined with N, P nutrition. This review is an effort to highlight the the
and potassium (K) fertilizers, mulching of residues role of essential mineral nutrients in improving
can improve yields by at least 1500 kg ha1 (Sun & agricultural WUE in crop plants.
Wang, 2001). Plastic film has also been widely used
to mulch the soil surface and promote crop growth
during early growth when temperatures are low. Macro-nutrients
Several methods of using plastic film have been
Nitrogen
adopted, including sowing wheat and rice through
holes in the plastic, sowing maize and wheat in rows Nitrogen is important in improving WUE and soil
in the furrow with plastic between the rows, and water use. Increased yields and WUE from added N
mulching two subsequent crops with the same were observed in several dry-land areas where crops
plastic film. Wang et al. (2004) conducted a field were grown on the same land for several years (Shan
experiment in a loess soil in central Shaanxi Province & Chen, 1993). Liu et al. (1998) reported maximum
to identify the effects of rainwater harvesting yield and highest WUE under optimum fertilizer
on WUE and yield of winter wheat. They used inputs of 90 kg N ha1 and 13 kg P ha 1 in the
ridge-furrow tillage, the ridge being mulched by semiarid field conditions of a loess hilly area in
plastic for rainwater harvesting in the furrows. Ningxia. Increased N fertilizer application was
They demonstrated that mulching significantly positively correlated with yield (correlation coeffi-
increased the harvesting of rainwater and signifi- cient, 0.96) and WUE (correlation coefficient, 0.89)
cantly increased yield. Biomass and grain yield in the of spring wheat. They also reported that N fertilizer
mulched plots were 39.5 and 28.9% higher than in applied in spring wheat improved the development
the corresponding treatments without mulching. of the root system and especially enhanced the root
 294 E.A. Waraich et al.

growth in the cultivated 0
20 cm soil layer. The molecules, it plays an important role in cell meta-
increased root system in the fertilized plants was able bolism. Absorption and utilization of nutrients like
to improve crop water use and nutrient absorption N by plants growing under water stress is very
and hence crop yield and WUE were increased. He critical for plant growth and productivity. Water
et al. (1999) conducted experiments to clarify the stress results in a depression in nutrient uptake
effects of water and N fertilizer and animal manures particularly of N which may contribute towards
on WUE of potatoes. The results showed that both reduced yield. In addition, although the crop may
N and water supply very significantly increased have roots penetrating the deeper and wetter parts of
WUE. Xu and Zhao (2001) reported that in north- the soil profile, nutrients concentrated in a dry
ern China, the crop WUE increased steadily over surface may be unavailable to plants (Wright, 1982).
10 years from 0.22 to 0.90 kg m3 due to technolo- Availability of N and consequently the response to
gical developments. The increase in WUE was mainly N applied as fertilizers or in other forms is closely
caused by the establishment of water conservation related to the ability of plant roots to absorb water
measures, soil improvement, the adoption of new crop from soil. When water inside the plant declines
varieties and the continuous use of increasing amounts below a threshold level, stomata close which causes
of N fertilizers. Fertilizers including nitrogen have a a decrease in transpiration resulting in a reduction in
great potential to further increase the efficiency of water transport through the plant. This in turn,
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water use, especially for crops that grow in autumn affects the roots’ ability to absorb water and
when the water supply is plentiful. nutrients as effectively as under normal transpira-
Possible mechanisms to enhance WUE by im- tion. Inhibition of nutrient uptake by plants in dry
proving N nutrition in crop plants are presented in soils also relates to the nutrient transport in soil by
Figure 1. mass flow and diffusion (Mackay & Barber, 1985;
Nitrogen, an important macronutrient, affects Seiffert et al., 1995), which may diminish nutrient
photosynthetic capacity of leaves by increasing availability at the root surface as well as cause a
stromal and thylakoid proteins in leaves (Makino decrease in mineralization of organically bound
et al., 1992; Bungard et al., 1997) and being an nutrients (Bloem et al., 1992; Walworth, 1992).
integral part of DNA, RNA, chlorophyll and other Severe drought may further decrease nutrient

Nitrogen (N)

– +
NO3 NH4

Increases Protein Increases Increases Membrane

Increases plant biomass
contents Membrane stability polarization

Increases stomatal and Increases SOD & Increases uptake of nutrients

Reduces ROS
Thylakoid proteins POD and water

Maintenance of chloroplast Increases Lipid Increases Stomatal

Membrane biosynthesis regulation

Reduces oxidative
damage Enhances Photosynthesis

Enhanced Water
Use Efficiency

Figure 1. Possible mechanisms through which nitrogen improves water use efficiency in crop plants. Please see text for abbreviations.
 Agricultural water use efficiency 295

transport to the root surface by inducing root of mild to moderate drought stress in soil-grown
shrinkage and which thus causes loss of soil
root plants.
contact (North & Nobel, 1997). The application of P fertilizer can improve plant
Inorganic fertilization has been reported to miti- growth considerably under drought conditions (Ack-
gate the adverse effects of water stress by increasing erson, 1985; Studer, 1993; Garg et al., 2004). The
the WUE of crop growth and development (Marschner, positive effects of P on plant growth under drought
1995; Payne et al., 1995; Raun & Johnson, 1999). have been attributed to an increase in stomatal
Under mild water stress low doses of N increased conductance (Brück et al., 2000), photosynthesis
grain yield in winter wheat, however under (Ackerson, 1985), higher cell-membrane stability,
severe water conditions high N application proved water relations (Sawwan et al., 2000) and water-use
to be detrimental (Nielsen & Halvorson, 1991). efficiency. Ajouri et al. (2004) reported that priming
Eghball and Maranville (1993) observed a non- seeds with solutions containing the limiting nutrients
significant interaction between drought and N under drought conditions (such as P and Zn) can
supply for root/shoot ratio, root length, root dry improve barley establishment. Smith (2002) sug-
weight, average root radius and nitrogen flux. Kathju gested that strategies for increasing nutrient uptake
et al. (1990) observed that when wheat plants were by overexpressing genes encoding for high-affinity P
grown under low (N0P0) and high (N80P80) fertility transporters are likely to be an important strategy in
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conditions and water stress was imposed at various the future, especially in light of the increasing
stages of a plant’s life cycle, increasing intensities of problems caused by P-deficient soils of the semi-
stress adversely affected leaf metabolism and plant arid tropics. Possible mechanisms to enhance the
performance. However, the performance of plants WUE by improving P nutrition in crop plants are
was better under high fertility conditions, at all presented in Figure 2.
stages, under different intensities of water stress.
Ashraf et al. (2001) reported that N application Potassium
minimizes the adverse effect of drought on dry
matter and grain yield in pearl millet. This indi- Potassium is an essential factor in protein synthesis,
cates that in dry land agriculture, where water is a glycolytic enzymes and photosynthesis; an osmoti-
limiting factor, fertilizer application to a reasonable cum mediating cell expansion and turgor-driven
extent can be considered for drought mitigation movements; and a competitor of Na  under saline
management. conditions (Marschner, 1995). Numerous studies
have shown that the application of K fertilizer
mitigates the adverse effects of drought on plant
Phosphorus growth (Andersen et al., 1992; Studer, 1993;
Phosphorus being a constituent of nucleic acids, Sangakkara et al., 2001). Potassium increases the
phospholipids, phosphor-proteins, dinucleotides and
adenosine triphosphate is required for processes Phosphorous (P)
including the storage and transfer of energy, photo-
synthesis, the regulation of some enzymes and the
transport of carbohydrates (Hu & Schmidhalter, Improves root Maintains high leaf
growth water potential
2001). Soils in arid areas are often calcareous and
have high pHs (e.g. those in Mediterranean regions).
In the semi-arid tropics, soils are often rich in Improves water and
Maintains cell turgidity
aluminium and iron oxides, and the pHs are low. nutrient uptake
Both of these soil types show a strong tendency for P
fixation (Oertli, 1991). It is generally accepted that
the uptake of P by crop plants is reduced in dry-soil Improves assimilation Increases stomatal
conductance
conditions (Pinkerton & Simpson, 1986). For ex-
ample, the translocation of P to the shoots is severely
restricted even under relatively mild drought stress
Increases photosynthetic
(Rasnick, 1970). However, Liebersbach et al. (2004) rate
reported that the large amount of molecular exu-
dates (i.e. mainly mucilage) from plants in dry soil Improves water use
efficiency
counteract the reduced mobility of P under such
conditions. Turner (1985) pointed out that P Figure 2. Possible mechanisms through which phosphorus im-
deficiency appears to be one of the earliest effects proves water use efficiency in crop plants.
 296 E.A. Waraich et al.

plant’s drought resistance through its functions in maintains the osmotic potential and turgor of the
stomatal regulation, osmoregulation, energy status, cells (Hsiao, 1973; Lindhauer, 1985) and regulates
charge balance, protein synthesis and homeostasis stomatal functioning under water stress conditions
(Beringer & Trolldenier, 1978; Marschner, 1995). It (Umar et al., 1993; Nandwal et al., 1988; Kant &
also maintains turgor pressure (Mengel & Arneke, Kafkafi, 2002), which is reflected in improved crop
1982) and reduces transpiration under drought yield under water limited conditions (Umar &
conditions (Andersen et al., 1992). In plants coping Bansal, 1997; Umar & Moinuddin, 2002). Besides,
with drought stress, the accumulation of K may be it takes part in many essential processes in plants
more important than the production of organic (Marschner, 1995) and enhances photosynthetic
solutes during the initial adjustment phase, because rate, plant growth and yield under stress conditions
osmotic adjustment through ion uptake like K is (Sharma et al., 1996; Tiwari et al., 1998; Egila et al.,
more energy efficient (Hsiao, 1973). Working with 2001; Umar & Moinuddin, 2002). The protective
wheat, Morgan (1992) showed that the lines dis- role of K in plants suffering from water stress has
playing high osmotic adjustments had a high accu- been attributed to the maintenance of a high pH in
mulation of K in their tissues. In these lines, K stroma and against the photo-oxidative damage to
accounted for about 78% of all osmotica. By chloroplasts (Cakmak, 1997).
contrast, amino acids, which were the only other
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important contributor, constituted only about 22%.

Calcium
A recent study by Ma et al. (2004) showed that K
accumulation in the expanding leaves in three Calcium plays a vital role in regulating many
genotypes of Brassica napus oilseeds accounted for physiological processes that influence both growth
about 25% of drought-induced changes in osmotic and responses to environmental stresses. Included
adjustment. Possible mechanisms to enhance WUE among these are water and solute movement as
by improving K nutrition in crop plants are pre- influenced through the effects of Ca2 on mem-
sented in Figure 3. Potassium nutrition increases brane structure and stomatal function, cell division
crop WUE by utilizing the soil moisture more and cell-wall synthesis, direct or signalling roles in
efficiently than in K-deficient plants. The positive systems involved in plant defence and repair of
effects of K on water use efficiency may be through damage from biotic and abiotic stress, and rates
promotion of root growth accompanied by a greater of respiratory metabolism and translocation
uptake of nutrients and water by plants (Rama Rao, (McLaughlin & Wimmer, 1999).
1986) and through the reduction of transpirational As for many other elements, the addition
water loss (Beringer & Trolldenier, 1978). Also, K of supplemental Ca2 has beneficial effects. For

Potassium
(K+)

Maintains High Enhances Root Decreases loss of Decreases transpiration

pH in Stroma growth soil moisture

Exploration of soil Increases Stomatal Increases retention

Reduces Photo-
water regulation of water in plants
oxidative damage

Maintains
Maintains Osmotic
chloroplast Maintains turgidity Enhances Photosynthesis
potential
Membrane

Enhanced Water use

efficiency

Figure 3. Possible mechanisms through which potassium improves water use efficiency in crop plants.
 Agricultural water use efficiency 297

instance, supplemental Ca2 in the growth medium ase and protein kinases (Marschner, 1995; Shaul,
increased the relative growth rate of barley under 2002). The role of Mg as the central atom of the
abiotic stress conditions (Cramer et al., 1990). chlorophyll (Chl) molecule is perhaps the best-
Similarly, Hu and Schmidhalter (1997) reported known function of Mg in plants in that it is
that although a biotic stress significantly reduced associated with the development of leaf chlorosis,
the Ca2 accumulation in wheat leaves, an increase typically interveinal, under Mg deficiency stress.
in the Ca2 concentration in the nutrient solution Depending on the Mg status of the plant, between
enhanced Ca2 accumulation. However, Hu and 6% and 35% of the total Mg may be bound in the
Schmidhalter (1998) later found no difference in chloroplasts, a higher proportion being associated
Ca2 accumulation between the control and stress with a lower Mg status (Scott & Robson, 1990).
treatments in growing leaves of wheat plants grown With the onset of Mg deficiency, there is firstly a
in soil. Supplemental Ca2 has also been reported to decline in the concentration of Chl b, which is
alleviate the adverse effects of stress on the germina- followed by that of Chl a (Hermans et al., 2004).
tion and vegetative growth of bean (Awada et al., An initial decrease in Chl concentration in Mg
1995). By contrast, various Ca2: Na  ratios had deficient plants has been ascribed to early accumula-
no significant effect on the uptake of Na  by rice tion of sugars in leaves, rather than to low levels of
(Yeo & Flowers, 1985). Mg (Hermans et al., 2004). According to Hermans
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In recent years, intracellular Ca2 has been found et al. (2004), high accumulation of sugars may
to regulate the responses of plants to drought and repress the expression of the cab2 gene that is
has also been implicated in the transduction of responsible for encoding Chl a and b protein.
drought stress signals in plants, which play an Magnesium deficiency symptoms vary between plant
essential role in osmoregulation under this condition species, but there are some similarities. As Mg is a
(Knight et al., 1997; Bartels & Sunkar, 2005). Key relatively mobile nutrient in plants, deficiency always
discoveries include drought-induced expression of appears first in the older leaves (Bergmann, 1992).
Ca2-dependent protein kinases (Urao et al., 1994) As discussed below in detail, development of Mg
and hyperosmotic shock-induction of putative deficiency symptoms is highly affected by light
Ca2-binding proteins (Ko & Lee, 1996). Sadiqov intensity. By increasing light intensity, plants under
et al. (2002) found that Ca2 participates in low Mg supply rapidly develop interveinal chlorosis
signalling mechanisms of drought-induced proline and reddish spots on the leaves (Marschner &
accumulation for osmotic adjustment. They sup- Cakmak, 1989). Moreover, for leaves with the
ported the findings of Knight et al. (1997), who same low concentration of Mg, development of leaf
indicated that this was due to the induction of a chlorosis and necrosis is significantly accentuated by
transcript for proline biosynthetic enzyme (P5CS) increasing light intensity (Marschner & Cakmak,
being inhibited in the presence of Ca2 channels in 1989). In crops, Mg deficiency is especially asso-
Arabidopsis (also shown in wheat and maize; Nayyar, ciated with strongly leached sandy acid soils with a
2003). low cation exchange capacity. One of the early
reactions of Mg deficiency stress in plants is its
impact on partitioning of dry matter between roots
Magnesium and sulphur
and shoots. An increasing body of evidence indicates
Little information is available on the role of Mg and that Mg plays a fundamental role in phloem export
S nutrition in enhancing water use efficiency of of photosynthates from the source to the sink organs,
plants. However, we expect that drought reduces and its deficiency results in dramatic increases in
both Mg and S uptake because Mg deficiency is accumulation of carbohydrates in the source leaves
occurring more frequently as cropping intensity (Cakmak et al., 1994a, 1994b; Marschner et al.,
increases and because the decrease in atmospheric 1996; Hermans et al., 2004). Reduced transport
S during the past decade has been associated with and hence accumulation of carbohydrates in Mg-
the appearance of S-deficiency symptoms (Scherer, deficient leaves cause alterations in photosynthetic
2001) and has had severe consequences for S carbon metabolism and restrict CO2 fixation. Im-
nutrition and crop production. pairment of the photosynthetic electron transport to
Magnesium is involved in numerous physiological CO2 through photosynthetic membranes may cause
and biochemical processes in plants affecting growth an accumulation of non-utilized electrons and ab-
and development. Magnesium is exceptional in sorbed energy. Under such conditions, the electrons
activating more enzymes than any other mineral and excitation energy not used in photosynthetic
nutrient (Epstein & Bloom, 2004). Some examples CO2 fixation is channelled to molecular O2, leading
of Mg-activated enzymes include ATPases, ribulose-1, to the generation of highly reactive O2 species
5-bisphosphate (RuBP) carboxylase, RNA polymer- (ROS) and consequently to damage of chloroplast
 298 E.A. Waraich et al.

Magnesium Boron
(Mg)
Boron is directly or indirectly involved in several
physiological and biochemical processes during plant
Increase root Increase root
uptake of nutrients growth and growth such as cell elongation, cell division, cell wall
and water surface biosynthesis, membrane function, nitrogen (N) me-
tabolism, leaf photosynthesis and uracil synthesis
(Marschner, 1995; Zhao & Oosterhuis, 2002).
Increase More uptake of
transport of water
Boron nutrition improves cell enlargement in
sucrose in root
growing tissues because of its structural role in the
cell wall (O’Neill et al., 2004). In higher plants, B
Reduce Carbohydrate
Increase in accumulation in
application induces changes in carbohydrate meta-
phloem export tissues bolism (Camacho-Cristobal & Gonzalez-Fontes,
1999), which in turn is responsible for the decrease
in concentration of phenolic compounds in leaves
Increases the
Under reduction
phloem export (Blevins & Lukaszewski, 1998; Camacho-Cristobal
of e-transport
et al., 2002). The low level of these phenolics oxidized
to derivatives such as quinones, reduce the produc-
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Increases tion of extremely reactive oxygen species (ROS) and

Reduces ROS
generation
photosynthesis cause reduction in peroxidative damage to vital
components of cell membrane such as lipids and
proteins, which eventually leads to repair of several
cellular functions (Cakmak & Romheld, 1997; El-
Enhances water Shintinawy, 1999; del Rio et al., 2003). Thus, B may
use efficiency
also protect plasma membrane against peroxidative
Figure 4. Possible mechanisms through which magnesium im-
damage by toxic O2 species. The role of B in
proves water use efficiency in crop plants. reproduction of higher plants is well established,
and its deficiency is responsible for creating male
constituents such as Chl and membrane lipids sterility and inducing floral abnormalities (Sharma,
(Marschner & Cakmak, 1989; Mittler, 2002; Asada, 2006). Apart from this, B deficiency is known to
2006). influence indirectly the activities of numerous en-
Possible mechanisms to enhance water use effi- zymes such as peroxidase, ribonuclease, indole acetic
ciency in crop plants by improving Mg nutrition are acid (IAA) oxidase, and polyphenol oxidase in several
presented in Figure 4. Magnesium increases the root crop plants. A key role for B has also been suggested
growth and root surface area which helps to increase in IAA metabolism and in regulation of lignin
biosynthesis and xylem differentiation (Marschner,
uptake of water and nutrients by roots and transport
1995). Boron nutrition improves sugar transport in
of sucrose from leaves to roots (Figure 4). Magne-
the plant body which helps to improve seed germina-
sium improves carbohydrate translocation by in-
tion and seed grain formation. This in turn improves
creasing phloem export and reduces ROS generation
yields and enhances water use efficiency (Figure 5).
and photo-oxidative damage to chloroplast under
Boron application also improves carbohydrate meta-
water-limited conditions. Maintenance of chloro- bolism and decreases the phenolic compounds in
plast structure by improving Mg nutrition enhances leaves. This in turn reduces the production of ROS
the photosynthetic rate under water stress which in and enhances the photosynthetic rate which results in
turn improves the water use efficiency. improved yield and water use efficiency (Figure 5).

Micro nutrients Zinc

The contributions of micro nutrients in enhancing Zinc is an essential nutrient for plant growth and
water use efficiency are not well-defined. Micronu- development. Crop productivity is limited due to its
trients help the macro nutrients in enhancing WUE deficiency (Irshad et al., 2004; Rashid & Ryan,
by activation of certain physiological, biochemical 2004). Zinc is involved in cellular functions such as
and metabolic processes within the plant body. The protein metabolism, photosynthetic carbon metabo-
role of micronutrients in drought alleviation is lism and indole acetic acid (IAA) metabolism
discussed below. (Marschner, 1995). Nutritional stress of zinc may
 Agricultural water use efficiency 299

Boron (B) Zinc (Zn)

Improve sugar Improve Carbohydrate Decreases activity

Increases IAA Increases auxin level Part of co-enzyme of membrane
transport metabolism
metabolism
bound-NADPH
oxidase
Improve seed Decrease phenolic Increases
Improves Causes bud
germination compounds in production of
regulation of Lignin swelling Decrease ROS
leaves tryptophane
biosynthesis generation

Increases auxin Precursor to

Improve seed and Reduces
Improves growth movement formation of Auxin Reduces photo-
grain formation production of ROS
oxidative damage

Enhances Enhances xylem Enhances

Improve yield differentiation root growth Enhances activities of
photosynthetic rate
SOD, POD, and CAT

Enhances water
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Enhances water use efficiency

use efficiency
Figure 6. Possible mechanisms through which zinc improves
Figure 5. Possible mechanisms through which boron improves water use efficiency in crop plants. Please see text for abbrevia-
water use efficiency in crop plants. tions.

affect plant water status, net CO2 assimilation rate, reduce (Marschner & Cakmak, 1989), while the
stomatal conductance and transpiration rate (Ben- activities of superoxide dismutase (SOD), perox-
Rouina et al., 2006). Zinc is a component of plant idase (POD) and catalase (CAT) enhance (Yu et al.,
carbonic anhydrase (CA) enzyme (Tobin, 1970) and 1998). Zinc plays an important role in lowering ROS
it is located in the chloroplast and cytoplasm in C3 generation and defending cells against ROS attack
plants (Okhi, 1976). A decline in CA activity has (Marschner, 1995; Cakmak, 2000).
been reported in plants due to Zn deficiency (Okhi,
1976). Earlier researchers (Nelson et al., 1969;
Silicon
Hatch & Slack, 1970) reported that CA may be
directly involved in photosynthesis, facilitating the Silicon (Si) has been regarded as an essential
diffusion of CO2 through the liquid phase of the cell element in a number of species of the Poaceae and
to the chloroplast. Sharma et al. (1995) observed that Cyperaceae, but it has not been possible to demon-
Zn deficiency caused decrease in CA activity, stoma- strate that it is essential to all higher plants because
tal aperture and transpiration in leaves of cauliflower. direct evidence is still lacking that it is part of
Possible mechanisms to minimize detrimental a molecule of an essential plant constituent or
effects of drought in crop plants by improving Zn metabolite (Epstein, 1994, 1999). Recently, the
nutrition are presented in Figure 6. Zinc is impor- role of Si in plant metabolism has received increasing
tant for its ability to influence auxin levels and has attention. Liang et al. (2003) strongly suggest that Si
long been known to be a co-enzyme for production may be involved in metabolic or physiological and/or
of tryptophan, a precursor to the formation of auxin structural activity in higher plants exposed to abiotic
(Bennett & Skoog, 2002). In most trees, the general and biotic stresses. It has been reported that Si
sequence of activity in early spring is auxin produc- increases plant tolerance to high manganese (Mn)
tion, bud swell, auxin movement, auxin-directed concentrations (Horst & Marschner, 1978), drought
transport, root growth, followed by phloem and (Lux et al., 2002), heavy metals (Neumann & zur
xylem differentiation (Seeley, 2006). As indicated Nieden, 2001) and plant resistance to pests and
above, normal auxin functions are likely to pathogens (Belanger et al., 2003; Richmond &
be disrupted in drought conditions. Maintaining Sussman, 2003). However, the use of Si and its
adequate hormone levels gives a competitive advan- relationships with plant growth and with mineral
tage to withstand adverse conditions of all kinds. In nutrition of plants under water stress have not been
another mechanism, Zn application reduces the studied in such detail.
activity of membrane-bound NADPH oxidase which The possible mechanisms to enhance WUE in
in turn decreases the generation of ROS (Cakmak & crop plants by improving silicon nutrition are pre-
Marschner, 1988) and photo-oxidation processes sented in Figure 7. Silicon enhances WUE by
 300 E.A. Waraich et al.

these nutrients in enhancing water use efficiency is

Silicon (Si)
described below.
Iron (Fe) is involved in the production of
chlorophyll. It is a component of many enzymes
Increases Improves Si Decreases
antioxidant accumulation in roots transpiration associated with energy transfer, nitrogen reduction
production and fixation, and lignin formation. Iron is associated
with sulphur in plants to form compounds that
Decreases Decreases apoplastic Increases catalyse other reactions. Drought-induced deficiency
ROS movement of retention of
generation heavy metal water in plants of Fe causes yellowing of leaves due to low levels of
chlorophyll. Leaf yellowing first appears on the
Decreases photo-
younger upper leaves in interveinal tissues. Severe
oxidative damage Improves Fe deficiencies cause leaves to turn completely
Reduces heavy stomatal
metal stress regulation
yellow or almost white, and then brown as leaves
die. Uptake of Fe decreases with increased soil pH,
Maintains and is adversely affected by high levels of available
chloroplast
membrane Reduces ROS Enhanced phosphorus, manganese and zinc in soils.
generation potosynthesis Manganese (Mn) is necessary in photosynthesis,
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nitrogen metabolism and to form other compounds

required for plant metabolism. Interveinal chlorosis
Enhances water
is a characteristic Mn-deficiency symptom. In very
use efficiency severe Mn deficiency cases, brown necrotic spots
appear on leaves, resulting in premature leaf drop.
Figure 7. Possible mechanisms through which silicon improves Delayed maturity is another deficiency symptom in
water use efficiency in crop plants. some species. Manganese has no direct role in
enhancing WUE. It can reduce the adverse effects
reducing the detrimental effects of drought (Epstein, of drought and improve WUE indirectly by enhan-
1999; Ma & Takahashi, 2002; Richmond & cing the photosynthetic rate, and nitrogen metabo-
Sussman, 2003; Ma, 2004). Silicon deposition in lism in the plant body. Manganese nutrition reduces
cell walls of roots reduces apoplastic bypass flow and interveinal chlorosis, brown necrotic spots on leaves
provides binding sites for metals, resulting in de- and reduces premature leaf drop.
creased uptake and translocation of toxic metals and Molybdenum (Mo) can reduce the adverse effects
reduces heavy metal stress injury to the roots. This of water stress and improve WUE indirectly due to
will improve the root activity and enhances the WUE its involvement in enzyme systems relating to nitro-
in plants. Deposition of Si in the culms, leaves and gen fixation by bacteria growing symbiotically with
hulls enhances the strength and rigidity of cell walls legumes, nitrogen metabolism, protein synthesis and
and decreases transpiration from the cuticle and, sulphur metabolism. It has a significant effect on
thus, increases the resistance to lodging and en- pollen formation, so fruit and grain formation are
hances WUE. Under water-limited conditions, affected by Mo nutrition in plants.
Si-alleviated effects have been associated with an Chloride (Cl) is a mobile anion in plants, most of
increase in antioxidant defence abilities (Liang et al., its functions relate to salt effects (stomatal opening)
2003; Zhu et al., 2004). However, this might be a and electrical charge balance in physiological func-
beneficial result of Si as opposed to a direct effect tions in plants. Chloride also indirectly affects plant
because it is unlikely that Si affects the activity of growth by stomatal regulation of water loss. Chloride
antioxidant enzymes. Silicon nutrition increases enhances WUE by reduction in wilting and restrict-
antioxidant production and reduces ROS generation ing highly branched root systems in cereal crops.
which in turn reduces photo-oxidative damage and The role of Cl in decreasing the incidence of various
maintains the integrity of chloroplast membrane. diseases in small grains is perhaps more important
This enhances the photosynthetic rate of the plants than its nutritional role from a practical viewpoint.
which in turn enhances productivity and water use
efficiency in plants.
Conclusions
Improving agricultural water use efficiency will
Other nutrients
continue to increase because of the demand for
The role of micro nutrients (iron, manganese, increased grain production. Greater yield per unit of
molybdenum and chloride) in enhancing water use water is one of the most important challenges in
efficiency is not well-defined. The indirect role of water-limited agriculture. In plants, drought can
 Agricultural water use efficiency 301

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