全 文 ：作物学报 ACTA AGRONOMICA SINICA 2013, 39(5): 761−766 http://www.chinacrops.org/zwxb/
ISSN 0496-3490; CODEN TSHPA9 E-mail: xbzw@chinajournal.net.cn

This work was supported by grants from the National Natural Science Foundation of China (Grant number: 30871447) and the Ministry of
Science and Technology of China (Grant number: 2011AA100501).
Corresponding author: MEI Xu-Rong, E-mail: meixr@ieda.org.cn; Tel & Fax: 86-10-82109333
Received(收稿日期): 2012-08-23; Accepted(接受日期): 2013-01-16; Published online(网络出版日期): 2013-02-19.
URL: http://www.cnki.net/kcms/detail/11.1809.S.20130219.1021.014.html
DOI: 10.3724/SP.J.1006.2013.00761
Improving Water Use Efficiency of Crops by Exploring Variety Differences
MEI Xu-Rong, ZHONG Xiu-Li, and LIU Xiao-Ying
Institute of Environment and Sustainable Development in Agriculture, Chinese Academy of Agricultural Sciences / State Engineering Laboratory of
Efficient Water Use and Disaster Mitigation for Crops / Key Laboratory for Dryland Agriculture, Ministry of Agriculture, Beijing 100081, China
Abstract: Improving water use efficiency (WUE) is considered to be an important measure for mitigating the conflict between
water resource crisis and sustainable crop production. In this review, variety differences in WUE at different time-space scales, the
scaling-up of WUE, and the association between WUE and yield are discussed. WUEintrinsic (WUEi) shows wide genotypic vari-
ability, in particular, under water deficit conditions. Variation in WUEi seems to be associated rather with variation in stomatal
conductance in cereals. WUEplant (WUEp) differed larger among genotypes under water deficit conditions, in contrast with the
smaller difference under well-watered conditions. Stomatal conductance is a determinant trait affecting WUEp based on the studies
performed up to date. Genotypes differ largely in stomatal conductance in response to water deficit. Scaling-up of WUE between
leaf level and field population level is limited by canopy and boundary layer resistances, partition of water use between soil
evaporation and plant transpiration and the internal allocation pattern of biomass. High WUEi associated with low stomatal con-
ductance can result in a considerable yield gain in a dry, stored-moisture, environment but it is likely to be disadvantageous in
terms of yield in more favorable growth environments.
Keywords: Water use efficiency; Yield; Water deficit; Variety difference
探讨品种间差异改良作物水分利用效率
梅旭荣 钟秀丽 刘晓英
中国农业科学院农业环境与可持续发展研究所 / 国家作物高效用水与抗灾减损工程实验室 / 农业部旱作节水农业重点开放实验室,
北京 100081
摘 要: 提高水分利用效率是缓解水资源危机实现作物可持续生产的重要策略。本文对叶片尺度的瞬时 WUE 和单
株尺度WUE的品种间差异, 瞬时WUE到田间尺度WUE的尺度转换, 以及瞬时WUE与产量之间的关系进行了讨论。
瞬时 WUE 具有较大的遗传变异性, 在亏水条件下品种间差异更显著。在禾谷类作物上, 气孔导度与瞬时 WUE 密切
相关。单株尺度 WUE在亏水条件下品种间差异显著, 足水条件下差异相对较小。气孔导度是影响单株尺度 WUE的
重要性状, 品种之间气孔对水分亏缺的敏感性差异较大。瞬时 WUE向田间尺度 WUE的尺度转换不仅受到冠层阻力
和边界层阻力的制约, 还受土壤蒸发与作物蒸腾比率以及同化物分配模式的影响。瞬时 WUE与产量的关系决定于环
境的水分条件, 在作物生长发育主要依靠土壤中储存水分的干旱条件下, 瞬时 WUE 高对获得高产有利。相反, 在水
分条件较适宜的地区, 高瞬时 WUE性状不利于高产。
关键词: 水分利用效率; 产量; 水分亏缺; 品种差异
Crop yield depends on the amount of water available
for growth, and the efficiency with which the water is
used in the water-limited environment [1-2]. In the regions
with deepening water-resource crisis, the only possible
way to ensure sustainable crop production is to signifi-
cantly improve water use efficiency of the crops. There is
considerable potential for raising WUE through im-
proved agronomic practices and also through breeding
for improved transpiration efficiency [3]. However, the
development of practicable and effective technologies for
increasing WUE is one of today’s greatest challenges[4].
Compared with agronomic water-saving methods, breed-
ing new crop varieties with a high yield and WUE has
the merits of being less investment intensive, greater up-
take, and sustainable efficiency to the grower, and there-
fore has a great promise for the future [5-6].
762 作 物 学 报 第 39卷

In recent decades, WUE had increased significantly in
modern cultivars [7], which is mainly attributed to efforts
in breeding for higher yield. However, the productive
varieties are usually the large water consumer. In a con-
text of declining water resources, further progresses in
WUE need to be achieved in which the water input is
reduced while the grain output remains high. WUE is the
result of an integration of many variables that affect plant
water consumption and yield formation throughout the
growing period. A comprehensive understanding of the
trait, including its determinant factors under specific
conditions, its responsiveness to the environment and the
trait’s contribution to yield, is required for its precision
evaluation and further improvement. Great efforts and
some progress have been made in developing means that
can predict crop varieties having high WUE and to dis-
tinguish ‘ideal’ crop varieties. This review focuses on
genotypic differences in WUE and its related parameters,
factors influencing WUE at different scales, the scal-
ing-up of WUE and the association between WUE and
yield. Stress will be laid on variations in the parameters
and the associations between them under water-deficit
conditions. Through the analysis, we will attempt to pro-
vide information that will be of practical use to breeders
and agronomic researchers.
1 The term “water use efficiency” defined
at different scales
WUE is a complex trait that is controlled by many
genes that are related to assimilate accumulation, water
consumption. The leaf WUE (WUEl), also called leaf
transpiration efficiency (TEl), is defined as the net CO2
assimilated by photosynthesis divided by the water tran-
spired in the same time period [8]. The intrinsic WUE
(WUEi) is the ratio between the net CO2 assimilated by
photosynthesis and stomatal conductance. WUEl and
WUEi are similar methodologically but the latter is not
influenced by the leaf-to-air vapor pressure deficit and
consequently, is used in comparative studies where dif-
ferent evaporative demands can be present [9-10].
Agronomists and crop physiologists define WUE rather
more from an integrative point of view, i.e. the accumu-
lated dry matter divided by the water used by the crop
over the same period [11]. Measurement of WUE is usu-
ally carried out at two different scales. In the studies in-
volved in variety differences, WUE is determined rather
more from individual plant scale, and described as WUE-
plant (WUEp) or TEplant (TEp), which is the ratio of eco-
nomic weight or biomass weight to the amount of water
transpired [12-13]. In some other studies in particular that
aiming at improving water management, WUE is meas-
ured at field population scale, and described as WUEfield
(WUEf, or water productivity). WUEf is defined as the
ratio of the total biomass or the economic product (e.g.
for cereals, just the grain) to the water consumed by
transpiration and evaporation from all surfaces [14]. In
calculation, the denominator for WUEf is both the water
transpired and the water evaporated from all surfaces, in
contrast of the WUEp, of which the denominator is the
water transpired only. Though WUE at different levels
can be related, the different spatial and temporal scales of
the concepts must always be carefully kept in mind.
2 Variation in water use efficiency of crop
varieties at multiple scales
2.1 Variation in WUEi of crop varieties under
different water conditions
WUEi and its related parameters such as net CO2 as-
similation rate and stomatal conductance can be easily
measured by gas exchange methods such as infrared gas
analyzer or porometry. To date, genotypic variation in
WUEi within a species has been reported for wheat [15],
cowpea (Vigna unguiculata) [16] and many other crops.
Genotypic differences may play important roles in the
WUEi variation found between different varieties of the
same species. In the case of some legumes (such as pea-
nut), variation in photosynthesis capacity accounts for
most of the variation in WUEi [17]. In many other cases,
however, variation in WUEi seems to be associated rather
with variation in stomatal conductance, as observed in
wheat [9,18-20], as found for common bean (Phaseolus
vulgaris Linn.) varieties by Ehleringer et al. [21]. The
variations in these parameters, coupled with other
physiological traits, provide a quick way to characterize
plant performance under water-deficit or under
well-watered conditions.
Net photosynthetic rate and stomatal conductance usu-
ally decrease under drought-stress conditions [22-23]. De-
crease in the net photosynthetic rate usually indicates a
stomatal closure factor in the presence of an increased
level of stress [23]. These parameters show a high degree
of variability under water deficit conditions [24-25]. Some
varieties strongly reduce stomatal conductance and in-
crease WUEi in response to relatively mild levels of
drought while some other varieties show smaller (<10%)
reductions in stomatal conductance under drought and no
rise at all in WUEi. The former were described as
“alarmist” varieties, the latter as “luxurious” ones. Ge-
netic variability in these traits can be an important source
of information to better appreciate the particular capaci-
ties of each variety to perform in arid environments [24],
thus studies on these should prove helpful for improving
the efficiency of breeding programs or for selecting
high-WUE varieties.
2.2 Variation in WUEp of crop varieties under
different water conditions
WUEp is related to stomatal regulation [25-26], but not
associated with stomatal frequency and size, according to
a recent study performed by Khazaei et al. [27]. WUEp
differences can also be related to differences in the pho-
第 5期 梅旭荣等: 探讨品种间差异改良作物水分利用效率 763


tosynthetic efficiency [28-30]. WUEp was observed to have
no significant correlation with antioxidants, proline, and
lipid peroxidation[31], and have poor association with
specific leaf area, SPAD chlorophyll meter readings and
carbon isotope discrimination (CID) [13,32]. Also, Bhat-
nagar-Mathur et al. [26] ascribed enhanced WUEp in the
transgenic events overexpressing DREB1A gene to a
lower stomatal conductance and an overall lower rate of
water loss per unit leaf area. These results imply that, for
WUEp, stomatal conductance is the important variable
found up to date.
High stomatal conductance confers both higher pho-
tosynthetic rate and higher transpiration rate, while low
stomatal conductance restricts both water loss and pho-
tosynthetic assimilation in the meanwhile. Due to this
fact, in some cases in particular under well-watered con-
ditions, though significant difference in biomass among
those varieties was shown, there was no significant dif-
ference in WUEp (e.g., Devi et al. [33]). However, geno-
types differ largely in sensitivity of stomata to soil drying
[33], and air drying [25,34]. Due to the fact that transpiration
rate declines more rapidly than photosynthetic rate with
decrease in stomatal conductance, genotypes with
stomatal conductance declining larger present higher
WUEp when encountering water deficit conditions, which
result in substantial difference in WUEp among geno-
types. Whether WUEp can be improved or reduced under
drought stress may depend on variety, stress extent, and
the growth stages exposed to stress. Under appropriate
drought stress, WUEp was observed to be improved, as
reported by some studies [35-37]. Ratnakumar et al. [38]
used a lysimeter system, a more reliable gravimetrical
method to assess WUEp over long period of time [39], to
assess the WUEp of nine peanut (Arachis hypogaea L.)
genotypes and presented varieties of higher along with
lower WUEp under water deficit than well-watered con-
ditions. Either way, variations in WUEp under drought
conditions can be ascribed most to differences in
stomatal conductance of different varieties in response to
water deficit, as discussed above.
2.3 Studies on WUEf provide more guidance for
agricultural practices
WUEf represent in practical terms the WUE of a vari-
ety in the field, and therefore deserve most interest.
However, the determination is difficult and laborious
requiring measurement of the water consumption of the
whole crop population. In recent years, some micromete-
orological methods such as the eddy-covariance method
and the Bowen ratio method have been adopted as the
best direct and scale-appropriate methods to assess WUEf
[40-42]. However, due to the larger land areas measured,
these methods are not suitable for comparative studies of
WUEf between different varieties which require only
relatively small plots. Till now, the number of compara-
tive studies performed on variety differences for WUEf is
far fewer than those for WUEl and WUEp. This is proba-
bly the lysimetric measurement developed recently offers
the possibility of measuring WUEf in plants that are cul-
tivated in field populations, where drainage is avoided
and where yield is measured [38-39]. Due to that WUEf is a
more complex trait determined by canopy characteristics,
environmental factors, in particular by agricultural prac-
tices, which all affect field evaportranspiration and crop
yield formation, studies on WUEf were conducted rather
more for a guidance of agricultural practices up to now.
3 The scaling-up of WUE
Both WUEi and WUEp are associated with stomatal
regulation and/or carboxylation efficiency [18-19,26-27].
WUEp can be closely associated with WUEi, as proved
by some earlier studies for wheat grown in pots [12,15,18].
WUEi and WUEf have implied associations by some
studies, which are either positively or negatively depend-
ing on the environment type. Van den Boogaard et al. [12]
compared the value of stomatal conductance and WUEf
of field-grown plants of eight wheat varieties and found
that, when plants were grown under rain-fed conditions,
the three varieties with the lowest stomatal conductances
had, on average, a 20% higher crop WUEf than those
varieties with higher stomatal conductances. No trend
was found under irrigated conditions. Similarly, experi-
ments carried out by Condon et al. [2] showed that the
variety with high WUEi also had a higher WUEf under
stored-moisture environments. In favorable environments,
however, the inverse trend was found. Bolger and Turner
[43] reported that, in Mediterranean annual pastures, sig-
nificant differences in WUEi observed in glasshouse ex-
periments seem to disappear under field conditions.
WUEi does translate to the field scale, as concluded by
Condon et al. [2], but is not always reflected in WUEf, in
particular in some more-favorable environments. This
also means that the use of carbon isotope discrimination,
which relates closely to WUEi, may not always be a good
screening method for WUE.
Scaling-up from WUEi to WUEf may be limited by a
number of factors. A reduction in stomatal conductance
does not necessarily lead to a significant decrease in
transpiration as might be expected. This could be due to
the fact that a reduced stomatal conductance is usually
associated with an increase in leaf temperature and an
increase in leaf-to-air vapor pressure difference, both of
which will counteract the decrease in leaf conductance
[28]. This counteracting effect will be higher in the case of
low leaf boundary conductance, low canopy aerodynamic
conductance, high levels of irradiance, high ambient
temperature and low relative humidity [44]. Moreover,
WUEp and WUEf are also influenced by biomass alloca-
tion patterns, i.e. the harvest index. Harvest index is con-
sidered to contribute majorly to variations in yield and is
regarded as an important trait of WUE in drought envi-
764 作 物 学 报 第 39卷

ronments and also in well-watered conditions. Further-
more, differences in crop WUEf could be masked by dif-
ferences in the partitioning of crop water use between
evaporation from the soil surface and transpiration from
plants such that differences in WUEf may become small
or insignificant [2]. WUEf is also influenced by a number
of aerial environment factors because the leaf-to-air va-
por pressure deficit determines the transpiration rate and
the same climatic factors also influence the evaporation
rate from the soil surface.
4 WUEi and yield
Some studies on the relationship of WUEl and yield
involve the measurement of carbon isotope discrimina-
tion (CID). CID is negatively correlated with WUEi,
which was proposed by Farquhar et al. [45] and thereafter
has been proved by a large number of studies in diverse
C3 plants [18,21,29,46-47]. Due to its convenience and despite
the expensive cost in determination, CID has become a
useful indicator of differences in WUEi. In recent dec-
ades, numerous studies have examined the association
between yield and CID. High correlations between CID
and grain yield have been reported, which were either
positive or negative. In Mediterranean-like environments,
where crop growth depends much on within-season rain-
fall, the correlations between CID and grain yield have
been mostly positive [48-53]. This contrasts with environ-
ments such as the northern Canadian prairies, where
wheat growth depends instead on stored-moisture, where
negative correlations between CID and aerial biomass
have been found [36,54-55]. Two new wheat genotypes,
which were higher yielding than the checks used, has
been released by selection for low leaf CID in wheat
breeding programs in Australia [56]. In addition to wheat
and barley (Hordeum vulgare Linn.), large genetic varia-
tions, and low genotype × environment interactions have
been reported for CID in field-grown sunflower (Helian-
thus annuus) [57], and cotton (Gossypium hirsutum L.)[58].
This implied the potential of CID as a reliable method
selecting varieties for high productivity. For more reli-
able selection, Anyia et al. [36] suggested that co-selection
for CID and other traits related to yield (e.g., harvest in-
dex) may be necessary for breeding programs aimed at
grain yield improvement.
The association between CID and yield indicated the
water condition-dependent relationship between WUEi
and yield. Van den Boogaard et al. [12] attributed the
negative relationship between WUEi and yield to the fact
that WUEi is limited by stomatal conductance whereas
the positive relationship to WUEi is likely limited by
photosynthetic capacity rather than by stomatal conduc-
tance. Using two winter wheat varieties with different
stomatal conductance, Condon et al. [2] found that high
WUEi associated with low stomatal conductance resulted
in a considerable yield gain in drier “stored-moisture”
environments. On the other hand, the “conservative”
gas-exchange associated with low stomatal conductance
can result in considerable yield reductions as well as in
less water use in favorable environments. In general, a
high WUEi would be disadvantageous in terms of yield
in favorable environments where crop yield depends on
current irrigation or on within-season rainfall. However,
in stored- moisture environments, improved WUEi is
likely to lead to consistent gains in crop yield. Thus,
WUE would appear to represent an important objective
for breeding programs for arid and semi-arid regions [59].
5 Conclusion
In the regions of water-resource shortage, the only
possible way to ensure sustainable crop production is to
improve water use efficiency of the crops. WUEi and the
related gas-exchange parameters present high genotypic
variation, in particular under water-deficit conditions.
The variations in these parameters provide some useful
information to characterize plant performance under wa-
ter-deficit or under well-watered conditions. For WUEp,
stomatal conductance is the important affecting factor
found up to date. Under well-watered conditions, no sig-
nificant difference in WUEp was shown in many cases.
However, genotypes differ largely in sensitivity of sto-
mata to soil and air drying. When encountering water
deficit conditions, genotypes with stomatal conductance
declining larger present higher WUEp, which result in
substantial difference in WUEp among genotypes. The
scaling-up of WUE still creates difficulties due to canopy
and boundary-layer resistances, the fractionation between
the water used for evaporation and that for transpiration,
the partitioning of assimilate to the grain compared with
that to the rest of the plant, as well as environmental fac-
tors. For WUE improvement, the associations of WUE or
its related traits with yield have to be taken account.
Whether the varieties with high WUEi associated with
low stomatal conductance can deliver higher yields than
those having low WUEi will depend greatly upon canopy
characteristics and environmental factors. High WUEi has
been proved by many studies to benefit high production
in water-limited environments, while be disadvantageous
in terms of yield in favorable environments. Though CID
has been proved to be a robust indicator of WUEi, the
correlation between CID and yield is environment type
dependent, indicating that co-selection for CID and other
traits related to yield may be necessary for breeding pro-
grams aimed at grain yield improvement.
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