els || water use efficiency of cultivated crops
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Water Use Efficiency ofCultivated CropsNader Katerji, INRA, Unite de Recherche Environnement et Grandes Cultures, Thiverval-
Grignon, France
Marcello Mastrorilli, Consiglio per la Ricerca e la sperimentazione in Agricoltura,
Agricultural Research Council – Research Unit for Agriculture in Dry Environments (CRA-SCA), Bari,
Italy
The concept of crop water use efficiency (WUE), deter-
mined by the ratio between the marketable yield and the
seasonal values of actual evapotranspiration, has become
a suitable tool for analysing the strategies that allow
attaining the best use of water in agriculture. Crop WUE is
easy to quantify from field measurements, but it is a
complex indicator because wide differences can be
observed for the WUE values of the same species culti-
vated under the same site.
In this article, the major causes of the large range of
WUE values are identified and analysed. The authors
demonstrate that the WUE variability can be ascribed
mainly to three factors: agro-techniques (water regime,
mineral supply and water quality), crop (species, varieties
and sensitivity of the growth stage to the stress) and
environment(climate,atmosphericpollution, soil texture
and climate change).
Understanding and taking into consideration the WUE
variability are primary conditions for advanced studies on
WUE. The paths for further research and management
programmes, allowing to valorise the water in agri-
culture, can be drawn from the analysis reported here.
Introduction
The irrigated cropping systems (differently from the rain-fed agriculture) are themainwater consumers of the planet.They utilised 64% of the total water withdrawals and 83%of the total consumptions in 2010. Water withdrawals andconsumptions increased six times on an average between1900 and 2010, nevertheless they could still increase by
18% between 2010 and 2025 (Roche and Zimmer, 2006).However, these global data mask a wide diversity at dif-ferent scales: regional, seasonal and interannual. Suchvariability is due mainly to the climate and to the raindistribution in particular, accompanied by the atmosphericevaporative demand, which is strictly linked to the airtemperature. The drier the climate, the more agriculturedepends on crop irrigation in order to obtain significantand regular productions and, as a consequence, waterwithdrawal does increase. Approximately 75% of thevegetal productions in water scarce regions come from theirrigated surfaces. In these regions the water allocation foragriculture use can attain 90% of the total withdrawals. Itis actually, the major constraint for other water uses(mainly municipal and industrial) that is at the base ofpeople’s well-being.Agricultural sector, especially in regions where water is
scarce, is currently faced with the challenge of newapproaches to water resource management that ensuresprotection of water resources and their integrity. Thisobjective can be attained by three types of strategies(Katerji et al., 2008):
. Save water by controlling water supply through betterdetermination of crop evapotranspiration, that is, waterrequirement (see the review by Rana and Katerji, 2000).
. Ameliorate the performance of irrigation systems sup-plying water to field through new techniques and meth-ods (see review of Pereira et al., 2002).
. Improve water use of cultivated crops.
The third strategy requires the identification and ana-lysis of the major causes acting on the water use of culti-vated crops. This topic is the objective of this article.
Methodology for Determination ofWater Use Efficiency (WUE)
Two approaches can be considered to determineWUE (seeKaterji et al., 2006b for extensive description).
1. The eco-physiological approach is based on the analysisat a given instant of the relationship between carbon
Advanced article
Article Contents
. Introduction
. Methodology for Determination of Water Use Efficiency
(WUE)
. The Concept of the Crop WUE
. A Review of the Value of WUE for Cultivated Crops
. Analysis of WUE
. Conclusion
Online posting date: 15th April 2014
eLS subject area: Plant Science
How to cite:Katerji, Nader; and Mastrorilli, Marcello (April 2014) Water UseEfficiency of Cultivated Crops. In: eLS. John Wiley & Sons, Ltd:
Chichester.
DOI: 10.1002/9780470015902.a0025268
eLS & 2014, John Wiley & Sons, Ltd. www.els.net 1
assimilation rate and transpiration rate per leaf unitarea. This approach helps reach the following results:
. To describe the processes (stomatal and nonstomatalconductance) determining water use efficiency at theleaf scale through theoretical approaches (Cowan,1982).
. To compare the leaf carbon assimilation and tran-spiration capacity of a species cultivated in relationwith soil water deficit conditions and the hormonalsigns from the root system (Damour et al., 2010).See also: Plant Response to Water-deficit Stress;Plant–Water Relations
. To underline the role of the breeding on leaf gasexchanges (Condon et al., 2004), and in particular, itsadvances onmolecular genetics approaches. See also:Abiotic Stress
The eco-physiological approachhelps in the understandingof global results obtained from the agronomical approach.It is evoked in the following paragraphs, which discuss thecauses of the following:
. the differences on crop water use values observedbetween C3 and C4 species,
. the effect of soil texture on crop water use values,
. the lack of a relationship between water use efficiencyand air pollution under water stress condition,
. the effects of the climate change on the water useefficiency.
However, it is not possible to calculate final crop yielddirectly fromcarbon assimilation rate per leaf unit area dueto the interference with many factors like respiration, leafgrowth, partition of assimilates flowering and pod setting.
2. The agronomical approach is based on crop waterconsumption and yield concepts. The time scale con-sidered is the whole vegetative cycle. They are key datato manage crop productions and for studying how tovalorise water in agriculture.
Ecophysiological and agronomic approaches are reallylinked, as showed byHsiao et al. (2007). The use ofwater toproduce food, according to these authors, is retained as aninterlinked chain of sequential efficiency steps. Their ana-lysis represents a relatively simple way to quantify theoverall water use in terms of the efficiency of each of thesteps, which can be divided according to engineering,agronomical and physiological processes.This article is aimed to analyse the WUE through the
agronomic approach. This approach refers to the sametime (the whole crop cycle) and space (the field plot) scalesretained by the three strategies, which have been identifiedin the introduction for setting up correct politics of waterresources management.
The Concept of the Crop WUE
Different concepts have been proposed and discussed tooptimise the crop water use (Pereira et al., 2002; Zwart and
Bsatiaanssen, 2004; Turner, 2004). The most known con-cept is the crop WUE (in kgm23), which results from theratio between yield (kgm22) and water consumption (i.e.actual evapotranspiration during cropping season(m3m22)). Yield is defined at the plot scale and it can beindicated by two parameters: biomass dry matter andmarketable yield.The marketable crop yield is a more interesting criterion
than dry matter, because in some species, like durumwheat, characterised by similar biomass, marketable yieldcan vary significantly as a result of genetic improvement(Katerji et al., 2005a,b).Moreover, it should be underlinedthat it is the marketable yield, and not the biomass, whichsettles on the economic value of any production.
A Review of the Value of WUE forCultivated Crops
Table 1 reports the crop WUE values for 16 species. Thisreview considers uniquely data from the Mediterraneanregion in order to minimise the role of the geographic andclimatic variability on WUE values (see section Climate).In addition, these values derive only from measured dataobtained from experimental setups on marketable yieldand on actual evapotranspiration.In general the crop productions seem highly costly in
water. Crops have to consume 1m3 (1000 l or kgs) of waterlost in air as evapotranspiration for producing 0.5 kgof grains from the broad bean or from the chickpea, 1 kgof grains from wheat, 6.5 kg of roots from sugar beet or oftubers from potato and 17 kg of table grape from thevineyards.The value of WUE of the species whose marketable
values are related to fresh weight (having a high watercontent in the vegetal tissues, as tomatoes and grape) arereasonably higher than the values observed for species withdry yield, like grain crops. For the latter ones, however,large differences exist between crops having C4 or C3metabolism. Corn (a species having the C4 metabolism) ischaracterised by aWUEvalue higher than that observed forcrops having the C3 metabolism, such as sunflower. Thesedifferences are explained by the analysis of the relationshipbetween carbonassimilation rate and stomatal conductanceobserved, at the leaf scale, on this two species (see Katerjiet al., 2006b). They can also be explained by seed compo-sition: corn contains essentially starch, whereas sunflowercontains 50% oil and 20% protein. The biosynthesis oflipids and protein is more expensive than starch.The large differences can be also observed between the
WUEvalues of the same species. These differences can existnot only in studies conducted in different countries, butalso in studies made at the same site. This can not only beseen for winter crops, like wheat, but also for summercrops, like corn (Table 1). This is the rationale of a series ofstudies aiming to analyse and understand the reason of thisvariability.
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Water Use Efficiency of Cultivated Crops
Analysis of WUE
The analysis of WUE at the crop scale consists of findingcorrelations between the observed values and a number of
parameters that are retained susceptible to explain theorigin of the WUE variability.The schema for analysing this variability was proposed
by Katerji et al. (2008). It consists in a synthesis of the
Table 1 Observed WUE values (kgm23) for species cultivated in the Mediterranean region
Grain crops
Wheat Syria 0.5–2.5 Oweis, 1997
Morocco 0.11–1.15 Corbeels et al., 1998
Morocco 0.32–1.06 Mrabet, 2002
Israel 0.6–1.60 Amir et al., 1991
Italy 1.02–1.20 van Hoorn et al., 1993
Italy 1.08–1.59 Katerji et al., 2005b
Turkey 1.33–1.45 Sezen and Yazar, 1996
Corn Turkey 1.65–2.15 Dagdelen et al., 2006
Turkey 0.22–1.25 Gencoglan and Yazar, 1999
Italy 1.35–1.80 Ben Nouna et al., 2000
Italy 0.82–1.17 Katerji et al., 1996
Lebanon 1.36–1.89 Karam et al., 2003
France 1.6 Marty et al., 1975
Spain 1.5–2.16 Fernandez et al., 1996
Barley Italy 1.46–2.78 Katerji et al., 2006b
Sunflower Italy 0.39–0.72 Katerji et al., 1996
France 0.6 Marty et al., 1975
Soybean Italy 0.47–0.77 Katerji et al., 2003
France 0.55 Marty et al., 1975
Lebanon 0.39–0.54 Karam et al., 2005
Sorghum Italy 0.67–1.59 Mastrorilli et al., 1995b
Broad bean Italy 0.86–1.37 Katerji et al., 2003
Italy 0.45–0.92 Katerji et al., 2005a
Syria 0.45–0.66 Oweis et al., 2005
Chickpea Italy 0.46–0.98 Katerji et al., 2005a
Syria 0.4–0.6 Oweis et al., 2004
Lentil Italy 0.36–2.09 Katerji et al., 2003
Syria 0.44–0.58 Oweis, 2004
Industrial crops
Cotton Turkey 0.61–0.72 Dagdelen et al., 2006
Turkey 0.50–0.74 Yazar et al., 1999
Israel 0.22–0.35 Saranga et al., 1998
Lebanon 0.8–1.3 Karam et al., 2006
Sweet sorghum Italy 3.69–4.20 (stalk) Mastrorilli et al., 1995a
Fresh yield crops
Potato Italy 16.2–18.5 Katerji et al., 2003
Sugar beet Italy 6.6–7.0 Katerji et al., 2003
Tomato Italy 4.4–8.3 Katerji et al., 2003
Italy 20 (globe fruit) Rana et al., 2001
Italy 22.2 (long fruit) Katerji and Rana, 2006
Fruit trees
Grapes Italy 16–18.1 Rana and Katerji, 2007
Clementine Italy 18.8 Rana et al., 2005
Source: Reproduced from Katerji et al. (2008).# Elsevier.
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Water Use Efficiency of Cultivated Crops
recent studies. According the schema shown in the Figure 1,the variability in WUE values derives from three mainfactors:
. Agro-techniques: water and fertiliser applied to cropsand analysed in terms of quantity and quality.
. Crop: differences between species, variety effects andphenological stage sensitivity to water constraints.
. Environment: soil and climate.
The last factor includes evaporative demand, atmosphericpollution and climatic changes.In reality, the different causes act together and indepen-
dently. For example, sunny days favour, at the same time,soil and atmospheric water deficits as well as the increasein air pollution. However, each identified cause will bediscussed separately in order to provide evidence for itsrole and its contribution to the variability of WUE values.
Agro-techniques factors
Water regime
The role of the water regime as a factor influencing WUEwas underlined byOweis (1997).Hemeasured the values ofwheat WUE in the semiarid environment of Syria forseveral years. Wheat was cultivated under irrigated andnonirrigated experimental conditions. This author noticedthat the average WUE of rain-fed wheat in Syria is0.5 kgm23, although with good management and favour-able rainfall amounts and distribution, this average couldbe increased to 1 kgm23. In fully irrigated areas with goodmanagement, the WUE was approximately 0.75 kgm23.
However, water used in supplemental irrigation conditionscan be much more efficient (WUE=2.5 kgm23). Thisextremely high WUE is mainly attributed to the effective-ness of a small amount of water in alleviating severemoisture stress during the most sensitive stages of cropgrowth.The averageWUE of rain-fed wheat observed in Syria is
close to the average (0.43 kgm23) observed in the MiddleEast–North Africa zone (de Fraiture andWichelns, 2007).The value is lower than those observed under the Medi-terranean climate of Australia. In fact, in south-westernAustralia, the average WUE of rain-fed wheat is approxi-mately 1 kgm23 (Sadras and Angus, 2006).
The previous observations, concerning the relationshipbetween WUE and water regime underlines two issues:
. the limit between WUE determined on irrigated andnonirrigated winter crops is not clear;
. the WUE of irrigated crops can present a large range ofvalues.
The generic term ‘irrigated crops’ or ‘rain-fed crops’ caninclude, in reality, extremely different situations of plantwater supply.Ben Nouna et al. (2000) pursued the lesson of Oweis
(1997) by quantifyingmore precisely the cropwater regimeduring the crop season. Three treatments, correspondingto three conditions of corn water status, monitored bypredawn leaf water potential measurements (the value ofthe leaf water potential determined before the sunrise),were identified: (1) IRRI, irrigated when predawn leafwater potential reached –0.3MPa. Such threshold value,
Parameters
Sources of variability
Environment Crop Agro-techniques
Variety
Water regime
Mineral nutrition
WUE
Soil texture
Phenological stage sensitivity
Climate
Air pollution
Climate changesWater quality
C3 and C4 species
Figure 1 Main factors affecting the WUE variability: Schema of analysis. Reproduced from Katerji et al. (2008). & Elsevier.
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Water Use Efficiency of Cultivated Crops
corresponds to the level at which maize leaf gas exchangesare affected bywater constraints. (2) STR1, irrigated, whenpredawn leaf water potential reached –0.6MPa. Thistreatment corresponded to moderate stress. (3) STR2,irrigated when predawn leaf water potential reached–1.2MPa. This treatment corresponded to severe stress.By using this field protocol for irrigation scheduling,
values of WUE calculated for maize decreased graduallywith the reduction of water supply, and it remained stablefrom one year to the next for the same water supply level(1.82 and 1.74 kgm23 under IRRI treatment, 1.61 and1.63 kgm23 for STR1 treatment and 1.35 and 1.53 forSTR2 treatment).The previous example provides evidence for the neces-
sity of reviewing studies on the relationship ‘water supply–WUE’ using a criterion appropriate to plant water status.The exact definition of the crop water status helps to dis-tinguish the limits between the different water regimes and,consequently, reduces the actual variability in WUEvalues.
Fertilisation supply
The action of mineral supply on WUE seems often to bestrictly related to water supply regime (Oweis, 1997; Oweiset al., 2000; Sadras, 2002, 2004; Zwart and Bastiaanssen,2004). As an example, Oweis (1997) noted that under rain-fed conditions, the rate of nitrogen fertiliser needed forwheat crop is not high, and that 50 kg ha21 is sufficient.However, with higher water supply, the crop responds tonitrogen up to 100 kg ha21 after which no benefit isobtained (Figure 2). This rate of N greatly improves WUE.It is also important tomaintain available phosphorus in thesoil so that the response to N and applied irrigation is notconstrained.The relationship ‘fertilisation–WUE’ for wheat growing
under the Mediterranean climate of Australia has been
discussed by Turner (2004) who simulated yield and eva-potranspiration on soils having different textures. Thesesimulations highlight the role of fertilisation technique inimproving WUE.
Water quality
Soil salinity corresponds to salt accumulation in the rootzone, leading to damage of cultivated plants. The soilsalinity can be characterised through the soil electricalconductivity (in dSm21) of the saturated soil paste extractsampled from the root zone. Themain causes of salinity arethe inappropriate management of irrigation and the use ofmarginal-quality water resources in agriculture, especiallyin water scarce environments.Soil salinity and soil water deficit have similar effects on
crops (Katerji et al., 2009, 2011). In fact, as salinity and soilwater deficit increase, the soil water availability decreases,and thismodifies plantwater status and gas exchange in theshort term, whereas growth and yield in the long term(Katerji et al., 2003).
In the frame of a long duration experiment carried on insouthern Italy, Katerji et al. (2008) analysed the con-sequences of using water of different salinity levels on theWUEof 10 species cultivated in lysimeter. The study showsa good correlation between species tolerance to soil salinityand their aptitude to maintain or to improve WUE whenirrigated with saline water. Based on the average values ofWUE obtained in clay and loam soils, a classification ismade and it led to the identification of two groups of spe-cies (Figure 3). The first includes wheat, sunflower, potato,maize and sugar beet. These species are tolerant to soilsalinity (see the review from Katerji et al., 2003) becausethey maintain, or slightly ameliorate, their WUE with anincrease in soil salinity. The second group includes tomato,lentil, broad bean and chickpea. These species are sensitiveto soil salinity, because their WUE is reduced with the
2.5
Rain-fed
1.0
1.5
2.0
WU
E (k
g m
−3)
Supplemental irrigation
0.0
0.5
0 50 100 150
Nitrogen application (kg ha−1)
Figure 2 Interaction between mineral supply and water supply regime on water use efficiency observed on wheat crop. Adapted from Oweis (1997) and
Katerji et al. (2008). & Elsevier.
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Water Use Efficiency of Cultivated Crops
increase in soil salinity. The previous observations areimportant for choosing the most appropriate species forsaline soils or for irrigation with saline water. Moreover,the soil salinity for certain species constitutes a cause ofvariability in the determination of WUE values.
Crop factors
This section discusses only the processes associated withcrop water supply responses.
Water stress deficit sensitivity at different cropgrowth stages
From the studies aiming to identify stress-sensitive phe-nological stages on species cultivated in theMediterraneanregion two interesting issues resulted:
. the critical stages correspond to major ones (e.g. flow-ering stage or fruit setting or assimilate transfer) in theelaboration of crop yield.
. during these critical stages amoderate water deficit leadsto a severe yield reduction and to a considerable varia-tion of WUE. In practice, this knowledge leads to theidentification of the conditions necessary for high
efficiency of water supply in relation to crop phenolo-gical stage sensitivity.
To define the sensitive stages, three methods are cur-rently used:
. by introducing soil water stress at a single phenologicalphase;
. by introducing at a single phenological phase a waterdeficit based on the ratio ‘actual evapotranspiration/maximal evapotranspiration’;
. by reducing the leaf water potential (through waterrationing) of plants at different phenological stages(Mastrorilli et al., 1995b, 1999).
Figure 4 and Table 2 provide an example of the methodbased on the field survey of the leaf-water applied to thegrain sorghum crop.When water stress was applied duringthe flowering stage (Figure 4a), the WUE value was one-third of that observed in the control and other treatments(Table 2). This reduction in WUE value is the result of thereduction in grain yield of 65% in respect to the controltreatment (Figure 4b).
The previous examples indicate that the irrigationcalendar, corresponding or not to sensitive phenological
0
30
60
90
0 2 4 6
WU
E (%
)
ECe (dS m−1)
(1) Wheat (1.04)
(2) Sunflower (0.21)
(3) Potato (18.5)
(4) Corn (0.98)
(5) Sugar-beet (7.0)
5)2)
1)
3)
4)
0
30
60
90
0 2 4 6
WU
E (%
)
ECe (dS m−1)
(6) Soybean (0.77) (7) Tomato (8.3)
(8) Broadbean (1.37) (9) Lentil (2.09)
(10) Chickpea (0.81)
10)7)
6)
8)
9)
0
30
60
90
0 2 4 6
WU
E (%
)
ECe (dS m−1)
(1) Wheat (1.04)
(2) Sunflower (0.21)
(3) Potato (18.5)
(4) Corn (0.98)
(5) Sugar-beet (7.0)
2)5)
1)
3)
4)
0
30
60
90
0 2 4 6ECe (dS m−1)
(6) Soybean (0.77) (7) Tomato (8.3)(9) Lentil (2.09) (10) Chickpea (0.81)(8) Broadbean (1.37)
10)7)
6)
8)
9)
Figure 3 Effects of soil salinity ECe on the water use efficiency of 10 species. ECe is soil electrical conductivity of the saturated soil paste extract.
Reproduced from Katerji et al. (2008). & Elsevier.
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Water Use Efficiency of Cultivated Crops
stages, has an important effect on yield, and consequently,it can be a cause of variability observed in WUE values.
Variety response to water stress
Many studies have been dedicated to the analysis of cropresponse to water stress caused by soil water deficit or soilsalinity in order to identify factors associated with planttolerance (see i.a. Munns, 2002) that may be taken intoconsideration for the creation of new resistant varieties(Ceccarelli et al., 2004).
Genetic improvements affecting the response of culti-vated species to water stress deficit have made importantprogresses in the past 20 years (Cattivelli et al., 2008). Thisprogress, however, has not involved salinity (Sharma andGoyal, 2003; Ashraf and Foolad, 2013).A significant number of recent studies has provided ele-
mentson the relationshipbetweenWUEandplant variety inthe presence and absence of saline stress for many Medi-terranean species. These studies concerned chickpea, broad-bean, durum wheat and barley (Katerji et al., 2005a, b,2006a, 2008).The study on durum wheat included the analysis of the
behaviour of varieties with different genetic origins.Experimental tests in greenhouses analysed yield andwater efficiency of seven varieties irrigated with three dif-ferent water qualities: fresh water, 4 dSm21, and 8 dSm21.
Irrigation was performed whenever soil water contentreached 30% of maximum water content value. Thisexperimental protocol avoided the exposure of plants tosoil water deficit.The seven studied varieties exhibited (Table 3) a large
range ofWUE, depending on irrigation water quality. Thefunctional classification of differentwater qualities allowedfor the identification of two varieties (Haurani andCham1)exhibiting the extreme behaviours at different salineconditions.The difference in WUE of these two varieties confirmed
the results obtained, under soil water deficit conditions, infield experiments (Annicchiarico and Pecetti, 2003) andthrough irrigation with different water qualities (Katerjiet al., 2005b). The WUE values obtained in this study arepresented in Table 4. The Cham1 variety had a higherWUEin comparison with Haurani when irrigated with freshwater, and it ameliorated its WUE when irrigated withsaline water. Under these conditions, both varieties pre-sentedWUE differences that reached approximately 40%.The previous example of wheat underlines two aspects:
. the importance of the choice of varieties resistant to saltor soil water deficit, in order to ameliorate WUE;
. the large variability in WUE, observed within the samespecies, can be attributed to a varietal response to thewater stress, caused either by soil water deficit or salinity.
30
50
70
90
Seed setting Grainformation
Control
Gra
in y
ield
(% o
f the
con
trol
)
(b)
**
n.s.
n.s.
Flowering−3.0
−2.5
−2.0
−1.5
−1.020 40 60 80
MPa
Days after emergence
(a)
Control
Flowering
Seed-setting
Grain formation
Figure 4 Grain sorghum crop: (a) predawn leaf-water potential (in MPa) measured during the crop cycle in 4 water treatments: Control (without any
water stress) + 3 temporary stress at a single phenological stage (flowering, seed-setting, grain formation); (b) grain yield (in % respect to the control)
measured from the 4 water treatments. Reproduced from Mastrorilli et al. (1995b). & Elsevier.
Table 2 Seasonal ET, water use efficiencies for grain (WUEg) and biomass (WUEb) for grain sorghum observed on the control
and temporary stressed treatments
Stress at ET (mm) WUEg (kgm23) WUEb (kgm
23)
Flowering (T1) 369 0.67 2.64
Seed-setting (T2) 360 1.59 4.01
Seed ripening (T3) 390 1.41 4.51
Never (Control) 420 1.51 4.85
Source: Reproduced fromMastrorilli et al. (1995b).# Elsevier.
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Water Use Efficiency of Cultivated Crops
Environmental factors
Climate
Climate plays a central role inWUE value. On the basis ofobservations made in dry environments (including LoessPlateau of China, Mediterranean region, North AmericaGreat Plains and south-eastern Australia), Sadras andAngus (2006) demonstrated that the variation in WUE ofwheat (ranging from 1 to 0.53 kgm23) was largelyaccounted for by reference evapotranspiration around theflowering stage. The daily evapotranspiration increases inrelation with the increase of air temperature and vapourpressure deficit (VPD).Higher WUE is achievable at lower VPD. The last
parameter represents the difference between the actual andmaximum pressure of water vapour in the air. The effectof VPDonWUEhas been the subject of previous analyses.A review conducted by Zwart and Bastiaanssen (2004) onWUE values observed for a winter crop (wheat) and threesummer crops (corn, rice and cotton), cultivated undersimilar experimental conditions between 10 and 40 degreeslatitude, is of considerable interest. These authors notedthat the VPD generally decreases when moving qawayfrom the equator and WUE is expected to increase withincreasing latitude. For example, the highest WUE valuesoccur between 30 and 40 degrees latitude where a two- tothreefold difference in WUE of wheat, rice and maize wasdetected when compared to areas between 10 and 20degrees latitude. Also, Rodriguez and Sadras (2007)reported a relationship existing between variations inWUE of wheat and latitude in eastern Australia.
The interannual climatic variability observed at the sameexperimental site affects the measured values of WUE aswell. Results of a study released by Katerji et al. (2010) insouthern Italy during a 25-year period, under experimentalneutrality (the same variety, sowing time, fertilisationsupply, soil texture, irrigation scheduling and water qual-ity), showed that the average value of WUE, observed onirrigated corn, is affectedby a standard error of 17%,whichis due exclusively to the climatic variability.
Atmospheric pollution
Some climatic conditions, high temperatures and solarlevels combinedwith stable airmasses and high emission ofair pollutants, favour the formation of secondary pollu-tants such as ozone (O3). Ozone concentration in theatmosphere is evaluated through indices, asAOT40.This isthe accumulated ozone over a threshold of 40 ppb duringan hour. When cumulated AOT40 during the growingseason exceeds a certain threshold, which depends on theozone sensitivity of each species, plant functions, mainlygas exchange, leaf growth, flower andpod setting, yield, areaffected (i.a. Morgan et al., 2003). However, the reportedresults did not provide any information about the rela-tionship between WUE and ozone exposure.One of the first studies on this subject was conducted by
Bou Jaoude et al. (2008a, b) in southern Italy on soybeancrops. Soybean was classified as a species sensitive to ozoneconcentration in atmosphere. WUE value, observed onwell-watered soybean subjected to high level of ozone dur-ing the vegetative cycle (cumulated AOT40=8500pbb.h),
Table 3 Classification of seven durum wheat varieties according to the water use efficiency (WUE) at three salinity levels of
irrigation water
WUE (kgm23)
ECiw=
0.9dSm
–1 V6 1.61 a
ECiw=
4.0dSm
–1 V5 1.68 a
ECiw=
8.0dSm
–1 V7 1.75 a
V7 1.59 a V7 1.61 a V5 1.69 a
V5 1.45 ab V6 1.49 a V6 1.41 b
V4 1.35 ab V4 1.26 b V4 1.40 b
V1 1.24 ab V2 1.21 b V2 1.04 c
V2 1.21 ab V3 1.09 bc V1 0.83 c
V3 1.16 b V1 0.93 c V3 0.78 c
Note: V1 Om Rabi-5; V2 Hagla; V3 Haurani; V4 Gidara-2; V5 Cham-1 (Waha); V6 Jennah Khetifa and V7 Belikh-2.Numbers followed by different letters are significantly different at the 5% level according to the Student–Neuman–Keuls test.Source: Reproduced from Katerji et al. (2008).# Elsevier.
Table 4 Interactive effects of three levels of soil salinity during the growing season and two durumwheat varieties (salt sensitive,
Haurani; salt tolerant, Cham-1) on seasonal actual evapotranspiration (ET), grain yield and water use efficiency (WUE)
Haurani Cham-1
Soil salinity (dSm21) 0.9 4.8 7 0.9 4.2 6.9
ET (m3m22) 0.72 0.64 0.60 0.68 0.60 0.55
Yield (kgm22) 0.83 0.69 0.66 0.97 0.96 0.87
WUE (kgm23) 1.15 1.08 1.10 1.43 1.58 1.59
Source: Reproduced from Katerji et al. (2005b).# Elsevier.
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Water Use Efficiency of Cultivated Crops
decreased by 30% in comparison with a control treatment(cumulated AOT40=0, obtained after air filtering). Con-versely, ozone had a not-significant effect on the WUEvalues observed on water-stressed soybean. Authorsdemonstrated that the absence of effect on water-stressedplants is due to stomatal closure, which reduces ozone fluxtowards leaves and thus its action on plant functions. Theseresults are the same as those observed in wheat (Khan andSoja, 2003), cotton (Heagle et al., 1988) and white clover(Fagnano and Merola, 2007).
Soil texture
The effect of soil texture, as a factor susceptible to mod-ifying WUE, was examined by Turner (2004). The authorreports that for wheat grown in Australia, WUE deter-mined on clay soil is lower thanWUEdetermined on sandysoil.Moreover, he highlights that the observed reduction isaffected by the level of nitrogen in the soil (see sectionFertilisation supply). However, these observations are notderived from measurements because they have beenobtained through simulations of yield, soil evaporationand plant transpiration.The study by Katerji and Mastrorilli (2009) on six spe-
cies (Table 5) is based onmeasurements of marketable yieldand seasonal actual evapotranspiration (ET) obtainedunder experimental neutrality (the same variety, sowingtime, fertilisation supply, irrigation scheduling and freshwater quality) in clay and loam soils.
TheWUE values observed in the loam soil (Table 5) weresystematically higher than those observed in clay. How-ever, the species can be classified into two groups. The firstgroup includes potato, corn, sunflower and sugar beet. Forthese species, the reduction in WUE in clay soil was sig-nificant and ranged from 22% to 25% with respect to theWUE values observed in the loam soil. This was due tosignificant reductions both in yield andETobserved in claysoil, with corn being the only exception. The reduction inWUE in this crop depended solely on a decrease in yield,whereas the differences in ET between the two soils werenot significant. The second group included soybean andtomato. For both species, the level of reduction in WUEobserved in clay soil was only 10%, which is not statisti-cally significant. In the case of soybean, there were nosignificant differences between the two soils for yield andET. Although in the case of tomato, only the variable yieldwas significantly lower in the clay soil.Convergent observations of stomatal conductance, leaf
water potential and actual evapotranspiration emphasisethat the water status of sugar beet plants growing in clay isless favourable than the water status of plants growing inloam (Katerji andMastrorilli, 2009). The more favourablecondition in loam soil has two origins:
. higher available water and easier soil water movements;
. higher root system capacity for water uptake.
The latter depends on root density and distribution,which in turn depends on the soil texture.
Table 5 Evapotranspiration (ET in mm), marketable yield (t ha21), WUE (kg m23) observed in different soil textures for the six
studied crops
Crop Clay Loam
Potato ET 363 415 a
Yield (tuber) 5.8 8.6 b
WUE 16.1 21.0 a
Corn ET 644 607 n.s.
Yield (grain) 0.55 0.68 b
WUE 0.87 1.13 a
Sunflower ET 1215 1450 a
Yield (grain) 0.22 0.35 b
WUE 0.18 0.24 a
Sugar beet ET 731 836 a
Yield (root) 4.47 6.56 b
WUE 6.11 7.85 b
Soybean ET 430 410 n.s.
Yield (grain) 0.31 0.33 n.s.
WUE 0.73 0.81 n.s.
Tomato ET 667 708 n.s.
Yield (fruit) 5.31 6.12 a
WUE 8.01 8.65 n.s.
ap40.05.
bp40.01.n.s.=not significant.Source: Reproduced from Katerji and Mastrorilli (2009).# Elsevier.
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Water Use Efficiency of Cultivated Crops
Climatic changes
Climatic change is due to the continuous rise of greenhousegases, primarily in the form of CO2. According to recentscientific reports, the effect of these variations on meanearth temperature levels are not yet known, especiallywhenthey are evaluated through general circulation models.These models were developed to take into considerationmany processes essential in determining future climate(radiation, convection, air mass circulation and turbulenceexchange with the surface), and they are susceptible, con-sequently, to important variability.In the analysis of the effect of environmental modifica-
tion on WUE, it is necessary to distinguish the effectsrelated toCO2 and temperature increases from thosewhichdepend on thewater regimes and their possible interactions(Katerji et al., 2008).
The rise in CO2 levels increase WUE values for the fol-lowing two reasons:
. The increase in photosynthesis results in a yield increase.C3 crops (wheat, rice and potato) increase yield up to30% when CO2 doubles. For C4 crops (corn and sor-ghum), however, the effect of CO2 levels on photo-synthesis is lower and final yield is not significantlyaffected.
. The reduction in stomatal conductance entails thereduction in transpiration of C3 and C4 plants. Formaize, with the doubling of CO2 level, transpiration isreduced by 15–17%. As a consequence, the WUE forbiomass increased by 24% (Bethenod et al., 2001).
Temperature increases modify WUE for two reasons:
. The reduction of crop cycle, which reduces water con-sumption (Campi et al., 2012).
. The increase of daily evapotranspiration due to theincrease in air in temperature and VPD (see sectionClimate).
The analysis of the interaction between temperature andCO2, associated with more or less important modificationson water regimes, is possible nowadays through the use ofcrop simulatingmodels that incorporate climatic scenariosexpected to occur in the future. Thus, these models givepredictions in relation to crop type,water amount, fertiliseruptake, and they test the strategies of water need andadaptation to climatic change.A recent example of such studies was performed using
two extreme climatic scenarios on corn grown under con-trasting water condition in southern Italy. This studyreported by Katerji et al. (2008) simulated the yield andwater use of corn through the Stics model (Brisson et al.,2003).In comparison with the 1984–2004 period, the corn
simulation in the 2070–2099 period, under both climaticscenarios, will reduce slightly the above-ground biomass.Conversely, grain yield is severely affected due to lowergrain weight. This yield component is reduced as a con-sequence of the shorter duration of the flowering-maturity
phase. Simulation also shows a reduction in seasonal eva-potranspiration. Finally, WUE values provided for thesimulations in the 2070–2099 period seem lower.However,due to a high variability, they do not differ significantlyfrom the values observed during the 1984–2004 period.For the C3 species results are different, as reported by
Asseng et al. (2004) for wheat, simulated in the Medi-terranean environment of western Australia. The increasein temperature andCO2 has a positive effect on grain yield,whereas evapotranspiration is reduced. According to theseobservations the WUE for wheat is expected to increase.However, after a more detailed simulation for differentAustralian sites and climatic scenarios, Ludwig andAsseng(2006) report that the predicted WUE increment varieswith seasonal rainfall distribution and differs significantlybetween soil types and locations.
Conclusion
The concept of crop WUE is easy to quantify from fieldmeasurements, but it should be retained as a complex indi-cator, because its intrinsic variability in relation with somefactors are identified and analysed in this article. Only if thecauses of such variability are known and taken into rightconsideration, the concept of WUE can represent an inter-esting tool for studying how to valorise the water in agri-culture. Studies on this topic should cover two pathways:
. The first type of studies are based essentially on fieldmeasurements and it consists of identifying and analys-ing the relation between the observed WUE values,obtained from different crops, and a number of para-meters that are retained as important to explain theorigin of the WUE variation and variability. The studyreported here is an example of this kind of approach.
. The second type of studies aim to highlight differentpotential strategies, based on agronomic and economicmanagements, in order to ameliorate crop WUE parti-cularly in areas where water resources are limited(Katerji et al., 2010; Andarzian et al., 2011; Garcia-Vilaand Fereres, 2012). For recommending solid watermanagement choices, each factor affecting crop WUEshould be previously identified and successively theirpotential interactions can be assessed. This objective isachievable through the correct use of crop models(Ritchie and Basso, 2008; Katerji et al., 2013). Never-theless, the choice of an operative model dependson its performance in correctly simulating, under con-trasting water stress conditions, the actual crop evapo-transpiration, biomass production and the finalharvestable yield. Many models can be generated tosatisfy the above exigencies, and these models havebecome available in the scientific literature over the pastthirty years. This is the case in some models: CERES(Jones and Kiniry, 1986), WOFOST (Van Diepen et al.,1989), EPIC (Jones et al., 1991), CropSyst (Stockle andNelson, 2000), STICS (Brisson et al., 2003), CRITERIA
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Water Use Efficiency of Cultivated Crops
(Marletto et al., 2007) and AquaCrop (Steduto et al.,2012).
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