Mutual physiological genetic mechanism of plant high water use efficiency and nutrition use efficiency.
ABSTRACT Water deficiency and lower fertilizer utilization efficiency are major constraints of productivity and yield stability. Improvements of crop water use efficiency (WUE) and nutrient use efficiency (NUE) is becoming an important objective in crop breeding. With the introduction of new physiological and biological approaches, we can better understand the mutual genetics mechanism of high use efficiency of water and nutrient. Much work has been done in past decades mainly including the interactions between different fertilizers and water influences on root characteristics and crop growth. Fertilizer quantity and form were regulated in order to improve crop WUE. The crop WUE and NUE shared the same increment tendency during evolution process; some genes associated with WUE and NUE have been precisely located and marked on the same chromosomes, some genes related to WUE and NUE have been cloned and transferred into wheat and rice and other plants, they can enhance water and nutrient use efficiency. The proteins transporting nutrient and water were identified such as some water channel proteins. The advance on the mechanism of higher water and nutrient use efficiency in crop was reviewed in this article, and it could provide some useful information for further research on WUE and NUE in crop.
- SourceAvailable from: cprl.ars.usda.gov[show abstract] [hide abstract]
ABSTRACT: In the Southern High Plains of the U.S.A., where water for irrigation is being depleted, drought-tolerant crops are extensively grown under limited irrigation where less water is applied than is required for potential evapotranspiration and maximum yield. This study was conducted (1) to determine the effects of plant water stress at various growth stages on growth and yield of soybeans [Glycine max (L.) Merr.] and (2) to assess the adaptability of the soybean plant to limited irrigation in the stressful climate of the Southern High Plains. The 3-year study was conducted on Pullman clay loam (fine, mixed, thermic Torrertic Paleustoll). Soybeans (‘Douglas’, indeterminate, maturity group IV) were grown with different irrigation treatments designed to subject the plants to water deficits at different growth stages. Stress initiated during R1 (early flowering) or R2 (full bloom) and extending to R3 (beginning pod development) reduced seed yields by 9–13%. But, when stress was extended to R4.5, yields were reduced by 46%. Stress beginning at R3 and extending to R4.5 reduced yields by 19%. Stress imposed at R5 and relieved at R6 reduced yields 15% in one year and 46% in a more stressful year. Stress imposed at R5 and extending to the end of the growing period (5 weeks) reduced yields by 45% in the less-stressful year and by 88% in the other. Stress throughout the last 3 weeks of the growing period (beginning at R6) reduced yields by 21 and 65%, respectively, in the two years. Water-use efficiency was not increased under limited irrigation. Soybeans are amenable to limited irrigation under the stressful climate of the Southern High Plains, but their vulnerability to drought stress during seed development complicates management. They are more suited for limited irrigation than is corn (Zea mays L.) but are less suited than are grain sorghum [Sorghum bicolor (L.) Moench], cotton (Gossypium hirsutum L.) or wheat (Triticum aestivum L.).Field Crops Research. 01/1987;
- [show abstract] [hide abstract]
ABSTRACT: The responses of gas exchange and water use efficiency to nitrogen nutrition for winter wheat were investigated under well-watered and drought conditions. The photosynthetic gas exchange parameters of winter wheat are remarkably improved by water and nitrogen nutrition and the regulative capability of nitrogen nutrition is influenced by water status. The effects of nitrogen nutrition on photosynthetic characteristics and on the limited factors to photosynthesis are not identical under different water status. Intrinsic water use efficiency (WUE(i)) of the plants at the high-N nutrition was decreased by a larger value than that of the plants in the low-N treatment due to a larger decrease in photosynthetic rate than in transpiration rate. Carbon isotope composition of plant material (delta(p)) is increased by the increase of drought intensity. The delta(p) at a given level of C(i)/C(a) is reduced by nitrogen deficiency. Leaf carbon isotope discrimination (Delta) is increased by the increase of nitrogen nutrition and decreased by the increase of drought intensity. Transpirational water use efficiency (WUE(t)) is negatively correlated with Delta in both nitrogen supply treatments and increased with the nitrogen supply.Environmental and Experimental Botany 11/2000; 44(2):141-149. · 2.58 Impact Factor
- Crop Science - CROP SCI. 01/1994; 34(1).
Colloids and Surfaces B: Biointerfaces 57 (2007) 1–7
Mutual physiological genetic mechanism of plant high
water use efficiency and nutrition use efficiency
Cao Hong-Xinga,f, Zhang Zheng-Bina,∗, Xu Pinga, Chu Li-Yed,
Shao Hong-Bob,c,d,e,∗∗, Lu Zhao-Huab,g, Liu Jun-Hongd
aCenter for Agricultural Resources Research, Institute of Genetics & Developmental Biology,
Chinese Academy of Sciences, Shijiazhuang 050021, China
bBinzhou University, Binzhou 256603, China
cQingdao Institute of Biomass Energy and Bioprocess Technology, Chinese Academy of Sciences, Qingdao 266071, China
dBiology Department, College of Molecular and Chemical Engineering, Qingdao University of Science and Technology, Qingdao 266042, China
eBioinformatics College, Chongqing University of Post & Telecom, Chongqing 400065, China
fGraduate University of the Chinese Academy of Sciences, Beijing 10039, China
gInstitute of Restoration Ecology, China University of Mining and Technology, Beijing 100083, China
Received 16 September 2006; received in revised form 15 November 2006; accepted 29 November 2006
Available online 6 December 2006
and biological approaches, we can better understand the mutual genetics mechanism of high use efficiency of water and nutrient. Much work has
Fertilizer quantity and form were regulated in order to improve crop WUE. The crop WUE and NUE shared the same increment tendency during
evolution process; some genes associated with WUE and NUE have been precisely located and marked on the same chromosomes, some genes
related to WUE and NUE have been cloned and transferred into wheat and rice and other plants, they can enhance water and nutrient use efficiency.
© 2006 Elsevier B.V. All rights reserved.
Keywords: Plant; WUE; NUE; Soil–plant root biointerfaces; Genetic mechanism; Crop breeding; Environmental effects; Developing directions
The advances in interaction of water and nutrient in plants.....................................................................
The evolution tendency of WUE and NUE in plants...........................................................................
The gene locating and molecular markers concerning WUE and NUE...........................................................
The gene cloning and protein function of WUE and NUE......................................................................
The transgenic relating to high WUE and NUE ...............................................................................
∗Corresponding author. Tel.: +86 311 85886648.
∗∗Correspondingauthorat:Center for Agricultural Resources Research, Institute of Genetic and Developmental Biology, Chinese Academy of Science,
Shijiazhuang,050021, China. Tel.: +86 311 85886648; fax: +86 311 85815093.
E-mail address: email@example.com (Z.-B. Zhang).
0927-7765/$ – see front matter © 2006 Elsevier B.V. All rights reserved.
H.-X. Cao et al. / Colloids and Surfaces B: Biointerfaces 57 (2007) 1–7
Water deficiency and low availability of nutrition often limit
crop growth and production potential in agricultural ecosys-
tems, because most of the crops are sensitive to water and
nutrient deficits in certain periods, particularly from flow-
ering to filling stages [1,2]. On the other hand, production
cost can be increased and environment is polluted by overuse
of fertilizer. So enhancing water use efficiency (WUE) and
nutrient use efficiency (NUE) in agriculture is very important
Water and nutrient uptake are two physiological processes
that interact with each other , which directly affects the dry
matter accumulation and distribution; the complex process has
a significantly positive effect on the economic properties of
crop and yield. Poor soil and improper nutrient management
often inhibit water effects, while irrigation could have a positive
effect only when fertilizer is properly applied [5–7]. There-
fore, it is imperative to understand the interaction mechanism
of nutrient and water for sustainable agriculture because water
and nutrient stress still exist in many areas around the world.
With the rapid development of molecular biology and genetics,
the uptake mechanism of nutrient and water has been studied by
ent uptake of plants in order to explain salinity accumulation
with time and plant growth [8–13]. Especially biotechnology
offers another avenue for modification of genomes by trans-
ferring one or more genes relating to improve WUE and NUE
by regulating their expression [14–16]. We can better study the
interaction of crop water and nutrient utilization from macro-
scopic to microcosmic domains. New theories and proofs will
be helpful for mapping quantitative trait loci (QTL) conferring
the drought resistance and nutrient deficiency, which will be
beneficial for developing cultivars with higher WUE and NUE
through marker-aided selection (MAS). The aim of this arti-
cle is to review the progresses of the interactions of high WUE
and NUE at physiological, genetic, molecular and biochemical
2. The advances in interaction of water and nutrient in
Water-use efficiency and nitrogen-use efficiency are impor-
tant indexes that affect the productivity of crops and had long
been intrigued many scientists to study. In the early 20th cen-
tury, Briggs and Shantz first studied the water requirement of
The definition of differ with the context in which it is consid-
ered, it is variously defined by agronomist, experts of plant
physiology, irrigation engineering and economists. In recent
study, WUE is briefly defined as the leaf WUE (leaf photosyn-
thetic rate per transpiration rate), whole plant WUE (the ratio
of aboveground biomass or dry matter per unit area, and water
use by crop); yield WUE (crop grain per unit area, to the tran-
spiration loss from the crop) . Plant nutrition was based
on the theory for mineral nutrition that was first brought for-
ward by the famous scientist Liebig . The study on NUE
could be traced back to 1920s when scientists put emphasis on
the process similar to the present nutrient revisited research,
which was also called the process of emigration or migration
From then on, there were many studies related to the interac-
tion between water and nutrient had been done in physiological
content. On the other hand, adequate fertilizer can improve the
water content of soil and crop WUE. In the 1980s, the famous
technique system named as “root growth was promoted by
phosphorus, more water was absorbed by more roots” was pro-
duced, which was one of the greatest achievements to improve
crop production in dry and semi-dry area in China . Many
researchers studied the variable of NUE under different soil
water content levels and the results indicate NUE has a big
difference under different water conditions [23,24]. Similarly,
plant WUE was affected by nutrient concentration and types
other important characters in plants. Different percentage of
P2O5in different crops under different water supplies, which
caused significant differences in dry weight . The interac-
tion among N supply, water potential, and air temperature was
exerted on leaf expansion in cotton plants (Gossypium hirsu-
tum L.) and has been indicated nitrogen deficiency considerably
increased the sensitivity of stomata to water stress. Stomatal
behavior may impart a stress avoidance type of drought resis-
tance to N-deficient plants . The seedling of Eucalyptus
kitsoniana Maiden and Eucalyptus globulus Labill were sub-
jected to two levels of water stress and two levels of nutrient
stress had been studied, the result showed water and nutri-
ent stress, applied separately or in combination, significantly
reduced leaf conductance, transpiration, photosynthesis and
midday water potential .
3. The evolution tendency of WUE and NUE in plants
Many studies have indicated that the capability of plants
to absorb water and nutrient has been improved in the evolu-
tion process. Modern cultivated wheat had been indicated to
have a higher WUE than their diploid and tetraploid ances-
tors in glasshouse experiments but they did not find consistent
differences between new and old varieties . The WUE
(grain yield/amount of water used) of modern cultivars was
higher than that of old varieties’ among nine Australian vari-
eties, because modern wheat cultivars with higher harvest index
. The leaf WUE, whole plant WUE and yield WUE was
increased with ploidy increased in wheat evolution from 2n to
is higher than that of dry land varieties, these results indicated
wheat WUE can be improved through breeding [31,32]. The
hydraulic and anatomical properties of different C3 and C4
woody species was investigated and found specific conductiv-
ity of stem xylem and leaf specific conductivity were lower in
C4 than in C3 shrubs, so the C4 shrubs exploits higher WUE
by altering xylem structure to enhance safety from cavitation
. Field studies were conducted at Hebei province from 1982
H.-X. Cao et al. / Colloids and Surfaces B: Biointerfaces 57 (2007) 1–7
to 2002 in China, the result indicated that crop yield improved
by 50% and water use efficiency (WUE) increased from 10 to
for maize .
The capability of plants’ water uptake was also found
gradually increased during the evolution process from diploid
to hexaploid and provided scientific evidences for WUE
breeding . The NUE evolution process also shows the
same increment tendency. Twenty genotypes of Triticum and
Aegilops wheat including diploid, tetraploid and hexaploid type
were investigated and the result showed that in the low-P
set, phosphorus efficiency increased as the yield and dry-
matter partitioning (harvest index) increased, with an order
of hexaploid>tetraploid>diploid. Low-P plants had simi-
lar grain nitrogen concentrations but lower nitrogen harvest
indexes than control plants . The changes of nitrogen
and phosphate use-efficiency of ten wheat evolution mate-
rials were studied, the experiment showed that nitrogen
use-efficiency gradually increased from diploid to tetraploid,
then changed little from tetraploid to hexaploid . A field
micro-plot experiment on the interrelationship between water
use efficiency and nitrogen use efficiency of different wheat
evolution materials at the whole plant level showed that both
water and nitrogen use efficiency increased gradually dur-
ing the long evolution process from diploid to hexaploid
Based on these findings and earlier evidences, we can draw
a conclusion that the WUE and NUE gradually increased dur-
ing crop evolution and agricultural production process. So high
WUE and NUE crop breeding should be paid more attention so
as to meet the need of agriculture production.
4. The gene locating and molecular markers concerning
WUE and NUE
factors. So it is difficult to evaluate the function of two traits
directly. A number of genetic maps for crop plants had been
established due to the advent of molecular markers. These maps
have been used to search for quantitative trait loci (QTL) for a
range of traits such as NUE and WUE. Great progress has been
achieved on the gene locating and molecular marker concerning
WUE and NUE.
We use wheat as an example, The flag leaf WUE of different
wheat genetic materials was measured and found that the order
of flag leaf WUE of different genomes is AA>BB>DD>RR.
The flag leaf WUE of A genome ditelosomie set is the high-
est. High WUE genes were located on 1AL, 2AL, 2AS and
7AS chromosome arms . The China spring-Egyptian Red
substitution lines was employed to make position of the genes
controlling leaf WUE and located genes on chromosomes 5A
and 5D .
The China spring-Cheyenne substitution lines was used to
study the genetic control of N-utilization and uptake in wheat,
the result showed that the gene control the nitrogen transfer
efficiency may be located on chromosomes 4B, 5B, 7B, and
association with K uptake and utilization efficiency was found
From the above said, we can see the gene location related to
WUE and NUE had been located on the same chromosomes,
which indicated the genetic of NUE and WUE may share com-
mon background in wheat.
In the molecular marker of NUE and WUE in wheat was
also had been studied. An F7recombinant inbred lines (RIL)
were used in a hydroponic experiment with phosphorus (P)
(0.25mmol/L) and deficient (0.025mmol/L) treatments and P
for SPUE under P deficient condition, chromosome 7A influ-
enced WPUE at both of the two P levels, that chromosome
5A was important for PUE under P sufficient environment
. The same recombinant inbred lines (RIL) population
(W7984×Opata85) was used to detect the location of WUE,
the shoot WUE was located on chromosomes 3B, chromosome
7A was vital for whole plant WUE, QTL related to whole plant
water consuming was mapped on chromosomes 5A . A
doubled haploid (DH) population of 150 lines, derived from
a cross between two Chinese common wheat varieties Hanx-
uan10 and Lumai14 was used in this study to map quantitative
at seedling stage under water stress and control conditions, the
result revealed that a total of 14 QTL of additive effect for the
7A, 1B, 3B and 3D . This doubled haploid (DH) population
was also used to map QTLs for N uptake and was located 1A,
1B, 5D, 7A .
These results of above research indicated that the same
genetic background on gene molecular markers of WUE and
NUE in wheat, which is very useful for molecular makers assis-
tance breeding to improve WUE and NUE in wheat.
Great progress has been made on the molecular marker of
WUE and NUE in other crops. For example, the chromosome
4 was found to control potential water use efficiency (13C) in
barley by combining the approaches of whole-shoot carbon dis-
genes had been proved on the chromosome 1R of rye (Secale
cereale L.) . The leaves of vegetative recombinant inbred
lines of maize was used to analyze physiological traits such as
(GS) activities, the result showed GS locus on chromosome 5
and this locus coincidences of QTLs for grain yield, GS, NR
activity and may be represent key . From above examples
we can find several traits are often related to one chromosome
and should be further studied.
These results could provide more information for plant com-
pare genomics of WUE and NUE, we could find more same
genetics background of WUE and NUE by plant compare
design breeding in future.
H.-X. Cao et al. / Colloids and Surfaces B: Biointerfaces 57 (2007) 1–7
5. The gene cloning and protein function of WUE and
Some proteins relating to water and nutrient uptake and
distribution have been identified and characterized. For exam-
ple, PM28A is an aquaporin and has been indicated that its
water channel activity is regulated by phosphorylation, and the
experiment results also suggest an active role for PM28A in
maintaining cellular water balance . The water and solute
transport characteristics of nodulin in 26 soybean was investi-
gated and the result showed that it is a multifunctional AQP that
confers water and glycerol, urea transport to the symbiosome
membrane, and likely plays a role in osmoregulation during
legume/rhizobia symbioses . Water channel protein RWC3
be possible regulate by GA and sucrose . The K+channel
decrease rice roots Lp by blocking water channels, K+content
of roots and leaves decreased after CsCl and TEACl treatment,
K+channel inhibitor increased the root and leave’s osmolarity
in rice . These experiments indicated the water and nutrient
transport has inner relation.
Many genes relating to NUE have been cloned due to the
introduction of new methods and technologies. The regula-
tory gene, areA mediating nitrogen metabolite repression in
ulatory gene mediating nitrogen metabolite repression . A
analysis of a high-affinity nitrate transporter in wheat (Triticum
tant role in high-affinity NO3-uptake . The gene designated
OsPTF1 has been cloned, in soil plot and field experiments
was indicated more than 20% increase in tiller number, pani-
cle weight, and phosphorus content was observed in transgenic
plants compared to wild-type plants at low-phosphorus levels
. In order to identify rice genes involved in nutrient par-
titioning, microarray experiments have been done to quantify
genomic scale gene expression, genes involving in nutrient par-
titioning specifically during grain filling, which will be used
to identify other co-regulated genes and DNA binding proteins
. The transcription factor Sfp1 was concluded that it could
integrate information from nutrient- and stress-responsive sig-
naling pathways to regulate ribosomal protein gene expression
From the above study results, we know some proteins and
ity and interact each other. The further study should emphasize
on to study the function of them. The further study should
emphasize on the study of their physiological and biological
6. The transgenic relating to high WUE and NUE
Drought resistance and high WUE has close relationship
in crop breeding. Some drought-resistant genes are used to
the HVA1 gene having higher water use efficiency values was
indicated and that growth characteristics were also improved
in transgenic wheat plants constitutively expressing the bar-
ley HVA1 gene in response to soil water deficits . The
ERECTA gene has been isolated from Arabidopsis and found
it can regulate transpiration efficiency and affect stomatal den-
sity, epidermal cell expansion, mesophyll cell proliferation and
method is a promising way for improving crop WUE.
in recent years. For example, the K channel gene was trans-
ferred to yeast, and the yeast could keep normal growth in the
low potassium medium . The K channel genes (AKT1 and
KAT1) were transferred to wheat by particle microprojection,
and found that the uptake, accumulation, utilization of potas-
sium of transgenic lines were improved to a certain degree .
Glutamine synthetase (GS) were showed play the central role in
plant nitrogen metabolism, transgenic plants containing trans-
ferred GS genes marked it is possible to improve crop nitrogen
use efficiency . The germplasm resources with high nutrient
use efficiency genes were identified in genetic analysis. Some
genes have been isolated which could remarkably improve the
nitrogen and potassium use efficiency in transgenic line. The
new varieties with high nitrogen and potassium use efficiency
will be cultivated in 8–10 years through transgenic techniques
and marker-assisted breeding approaches .
TheNUEis studied from
varieties→chromosome location→molecule marker→gene
cloning→analysis of gene function. But the research on WUE
was only from the differences among varieties→chromosome
remains unsolved. With the rapid progresses of molecular biol-
ogy and transgenic techniques, high water and nutrient use
efficiency genes may be cloned and transferred into plant at
one time, so that water and nutrient utilization efficiency will be
ing and they will play a vital role in improving the production
in the future. But water deficits and lower NUE are the most
overriding constraints for crop production and they often as the
been further studied [66–71]. From overview on crop breed-
ing history, every breakthrough needs germplasm innovation
and with molecular breeding so that excellent trait is produced
and reached. It is clear that plant molecular biology and plant
biotechnology are important technologies in increasing WUE
and NUE [72–76]. In order to develop crop cultivars suitable
for water and nutrient scarce environments, firstly we can select
excellent germplasm resources and identify the critical genes
relating to nutrient and water utilization in main crop plants.
Secondly, mapping QTLs conferring NUE and WUE and the
implementation of DNA markers in plant breeding programs
using marker assisted selection (MAS) as the main approach.
Thirdly, genetic engineering has become an important practice
H.-X. Cao et al. / Colloids and Surfaces B: Biointerfaces 57 (2007) 1–7
in agricultural biotechnology, many crops were cultivated by
this way and it also could be used in this aspect.
The new breakthrough in water saving would be attained
only by improving plant WUE by organisms themselves .
iological factors but modified by the interactions of plant with
environmental variables . So improving NUE and WUE of
plant is a complex issue which requires a mutual understanding
and an interdisciplinary approach because they are influenced
by many single factors. In recent years, many new methods
appeared providing powerful tool for genetic improvement of
WUE and NUE in crops, such as T-DNA insertions and dele-
tion mutants are currently used as a tool to identify phenotypic
mutants for characterizing relevant genes at the molecular level
and microarray-based expression profiling are used to isolate
and identify candidate genes. Gene-network estimation method
to facilitate analysis of gene interactions can serve as special-
ized tools in bioinformatics, which open new possibilities for
the development of new cultivars with higher WUE and NUE
[79–83]. In the future, we believe that more and more genes
directly controlling WUE and NUE will be cloned and trans-
ferred into the main crops. International collaborations between
biologists and breeders are important for breeding of high NUE
and NUE crops, and public acceptance is also important for the
success of transgenic plants.
From the above all information, the full physiological and
molecular relevance of NUE and WUE only in the initial stage,
able, valuable information for genetic breeding, the regulatory
mechanisms of response to water and nutrient stress is largely
remain unclear, there is still a long way to go.
We are grateful for the joint support from National Nat-
ural Science Foundation of China (30270821), Important
Knowledge Innovation Item of Chinese Academy of Sciences
(KSCXZ-SW-327), National 863 Water saving of Important
Item (2006AA100201) (Zhang, Z.B.). Qingdao University of
of Sciences, National Science & Technology Supporting Pro-
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