Agricultural Water Management 159 (2015) 6676

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Agricultural Water Management

journal homepage: www.elsevier.com/locate/agwat

Effect of partial root zone drying and decit irrigation on nitrogen and
phosphorus uptake in potato
Caixia Liu a, , Gitte H. Rubk a , Fulai Liu b , Mathias N. Andersen a
a
Department of Agroecology, Faculty of Science and Technology, Aarhus University, Blichers All 20, 8830 Tjele, Denmark
b
Department of Plant and Environmental Sciences, Faculty of Science, University of Copenhagen, Hjbakkegrd All 13, 2630 Tstrup, Denmark

a r t i c l e i n f o a b s t r a c t

Article history: Better understanding of the effects of decit irrigation regimes on phosphorus (P) and nitrogen (N) uptake
Received 22 December 2014 dynamics is necessary for sustainable water, P and N management. The effects of full (FI), decit (DI) and
Received in revised form 20 May 2015 partial root-zone drying (PRD) irrigation on potato P and N uptake with P fertilization (P1) or without
Accepted 22 May 2015
(P0) were investigated in two split-root pot experiments in a soil with low plant available P. Under FI, the
plants were irrigated to pot water holding capacity while under DI and PRD, 70% of the water amount of
Keywords:
FI was applied on either both or one side of the pots, respectively. During potato growth, plant P uptake
Water saving irrigation
increased while P concentration decreased at P1 and was almost constant at P0. PRD and DI reduced
Phosphorus and nitrogen uptake
Water use efciency
plants P uptake to a same extent, ca. 22% compared to FI at P1, while at P0, plants P uptake was similar
Drying and rewetting cycles for the three irrigation treatments. Soil P transport to the root surface by diffusion was similar under DI
Immobilization and PRD. DI treatments had higher soil microbial biomass P, water soluble P, root biomass and leaf water
potential than PRD treatments, while PRD treatments had higher plant N:P ratios than DI treatments
and higher root secretion of acid phosphatases that may have compensated for the lower level of water
soluble P. N was immobilized in soil in all the treatments. Plant N uptake under PRD was higher than DI
at both P levels, which could be explained by the higher microbial biomass and N-immobilization under
DI. In conclusion, when same amount of water was used, PRD was superior to DI in terms of N uptake,
but not P uptake. Challenges remain how to maintain crop yield and P uptake under reduced irrigation
regimes. Utilization of water and N fertilizer was low when the soil was decient in P.
2015 Elsevier B.V. All rights reserved.

1. Introduction
The increasing worldwide competition for fresh water resources
requires better management of irrigation to improve water use ef-
ciency (WUE). Decit irrigation (DI) and alternate partial root-zone
drying irrigation (PRD) are two water saving irrigation regimes,
which can increase WUE of potatoes (Wang et al., 2009) even with-
out reducing yield (Shahnazari et al., 2007). Under DI, the crop
receives irrigation water amounts slightly less than actual evap-
otranspiration but the resulting mild stress has minimal effects on
yield (English and Raja, 1996). PRD is a special form of DI where only
Abbreviations: P, phosphorus; N, nitrogen; FI, full irrigation; DI, decit irrigation;
PRD, partial root-zone drying irrigation; WUE, water use efciency; WSP, water sol-
uble phosphorus; P1, treatments with phosphorus fertilizer; P0, treatments without
phosphorus fertilizer; MBP, microbial biomass P; SNmin , soil mineral N balance;
APM, soil acid phosphomonoesterase activity.
Corresponding author at: Aarhus University, Blichers All 20, 8830 Tjele,
Denmark. Tel.: +45 87154761.
E-mail address: caixialiu21@hotmail.com (C. Liu).
half of the root zone is irrigated while the second half is exposed to
soil drying to a predetermined level before switching the irrigation
to this second half. To maximize potato yield, not only sufcient P
but also sufcient water need to be available (White et al., 2005a).
Since DI and PRD reduce the soil moisture content, the tortuosity
of the diffusion pathway for nutrients increase (Gahoonia et al.,
1994) and the transport of P from soil to root is reduced (McBeath
et al., 2012). Compared with DI, PRD increases the nitrogen (N)
availability by allegedly inducing greater microbial activity and
mineralization of organic N, which resulted in improved N uptake
in tomato (Wang et al., 2010a,b, 2012, 2013) and potato (Shahnazari
et al., 2008; Wang et al., 2009).
Potato is the fourth most important global crop by volume (FAO,
2007). Potato has a shallow root system (Vos and Haverkort, 2007),
is generally drought sensitive (Yuan et al., 2003), and has a high
phosphorus (P) requirement (Westermann, 2005). P is an essential
element for all plants and rock phosphate from which P fertiliz-
ers are derived is a vital non-renewable resource, which may be
depleted in a relatively short time span (Elser and Bennett, 2011).
However, plant P uptake of freshly applied fertilizer P is normally

http://dx.doi.org/10.1016/j.agwat.2015.05.021
0378-3774/ 2015 Elsevier B.V. All rights reserved.
 C. Liu et al. / Agricultural Water Management 159 (2015) 6676
modest since P tends to bind strongly to soil constituents e.g. in
Oxisols (Shamshuddin and Anda, 2012); with time, the immedi-
ate availability of added P fertilizer declines due to a combination
of abiotic and biotic processes (Park et al., 2004; Frossard et al.,
2000). Due to low P concentration in the soil solution, movement
of P in soil is dominated by diffusion, and a large P concentration
gradient occurs across the rhizosphere between the bulk soil and
the root surface (Tinker and Nye, 2000). At low soil P availabil-
ity plant adaptations to enhance P uptake are induced, including
increased secretion of phosphatase (Bargaz et al., 2012), exudation
of organic acids (Liao et al., 2006), greater root and root hair growth
(Balemi and Schenk, 2009; Lynch and Brown, 2001) and enhanced
expression of P transporters (Liu et al., 2010).
There is some evidence that irrigation management can also
inuence the use of organic P and the uptake of P (Wang and
Zhang, 2008, 2010, 2012; Wang et al., 2012). P interacts positively
with N uptake and plant growth (Fageria, 2001), so an improved
N uptake might be accompanied by an improved P uptake under
PRD. Although several studies on P uptake in potato (Alvarez et al.,
2001) and tomato (Wang et al., 2012) have been conducted, a thor-
ough study of how PRD and DI affect P availability and dynamic P
uptake in situations with scarce water and P resources is lacking.
The main objective of this study was therefore to investigate the
effect of FI, DI and PRD irrigation on P and N uptake in situations
with or without P fertilization.
2. Materials and methods
2.1. Experimental materials and design
Two pot experiments were carried out in a climate-controlled
greenhouse at Aarhus University Research Centre Foulum,
Denmark using cylindrical pots with a volume of 10 L (16 cm outer
diameter and 50 cm deep), divided into two vertical compartments
by a diametric plastic sheet. A 5 cm wide and 10 cm high piece was
cut and removed from the middle of the sheet at the top where the
seed tuber was going to be planted. Thus water exchange between
the two compartments was prevented. The bottom of the pots was
covered by 1.5 mm nylon mesh.
For the rst experiment (Exp1), the soil was collected from the
experimental farm of Foulumgaard also at Research Centre Foulum,
and was sandy loam, with 68% sand, 24.3% silt and 7.7% clay, total
C 16.3 g Kg1 , total N 1.5 g Kg1 , Olsen P 22 mg kg1 , water solu-
ble phosphorus (P) (WSP) 6.3 mg kg1 , mineral N:(NH4 + and NO3 )
7.4 mg kg1 , pH 5.4. The eld capacity and wilting point of the soil
at pF = 2.0 and pF = 4.2 were 25.3% (Vol.) and 6.7% (Vol.), respec-
tively. The water retention curve for the soil is shown in Fig. 1. The
soil was dried under a roof to a water content of ca. 12% (v/v) and
sieved through a 1 cm mesh sized sieve. Each pot was lled to a
dry bulk density of 1.3 g cm3 corresponding to a total of 11.17 kg
dry soil. The soil was divided into a subsoil layer with a depth of
24 cm that was unfertilized and a topsoil layer with depth of 20 cm
that was mixed with fertilizer before packing. In the second exper-
iment (Exp2), the sandy loam soil was collected from the same site
as Exp1 with Olsen P 22 mg kg1 , WSP 8.7 mg kg1 , pH 6.2. The soil
was air dried and sieved (1 cm mesh) and heated in an oven two
times for 24 h at 85 C with an interval of 48 h at room tempera-
ture to eliminate living microorganisms. Sterilized soil had Olsen
P 26 mg kg1 , WSP 12 mg kg1 , pH 6.0. Pots were pre-sterilized by
washing with 2% (w/v) NaClO and rinsed with tap water. Each pot
was lled with 10.84 kg dry soil to a dry bulk density of 1.3 g cm3
with topsoil and subsoil layer of 22 cm each.
Seed potatoes (Solanum tuberosum L. cv. Folva, size: 5580 g)
were pre-germinated. When sprouts emerged, one seed potato
was planted in the middle of each pot with all except one sprout
67
Fig. 1. Soil water retention curve for the experimental soil sampled from Ap-horizon
030 cm at Foulumgaard.
removed from the tuber. In Exp2, potato tubers were surface
sterilized with 0.5% (v/v) H2 O2 for 1 min and rinsed thoroughly
with distilled H2 O before the sprouting. Exp1 was conducted from
November 2012 to January 2013 in the greenhouse. Exp2 was car-
ried out from August to middle of October 2013 and the pots were
initially placed outside under a transparent roof for 20 days and
then moved into the greenhouse. The conditions of the green-
house in both experiments were set as: 20 C/10 C 2 C day/night
air temperature, 16 h:8 h L:D photoperiod and >500 mol m2 s1
photosynthetic active radiation (PAR) supplied by sunlight plus
metal-halide lamps. Pots were randomized on two tables in the
greenhouse.
In Exp1, two P fertilization levels were created: P1 in which the
20 cm topsoil layer of each pot was supplemented with compound
fertilizer consisting of 3.1 g N (47.73% NH4 -N, 52.27% NO3 -N), 2.57 g
K, 0.63 g S and 0.8 g P (CaHPO4 2H2 O); the control (P0) had the
same amounts of N, K, S but no P. The calculation of fertilizer
rates per plant was based on an assumed eld N application rate of
150 kg N ha1 and P rate of 37 kg P ha1 and further assuming that
potatoes are planted at 40,000 plants ha1 (Haverkort, 1982). The
packed soil had a water content of 24.4% (v/v) at pot water holding
capacity measured in situ after 2 days of drainage of the pots placed
on a moist, naturally drained outdoor soil surface and 5.18% (v/v) at
permanent wilting point as measured in a pressure plate apparatus
(Fig. 1). In Exp2, P1 was supplemented with 2.6 g N, 2.8 g K, 1.8 g S
and 0.6 g P mixed into the 22 cm topsoil, while the control (P0) had
no P but otherwise the same dressing. Fertilizers used were pure
chemicals: NH4 NO3 , KH2 PO4 , K2 SO4 , MgSO4 7H2 O. The packed soil
had a water content of 23.8% (v/v) at pot water holding capacity.
Soil water content was monitored by a time domain reectome-
ter (TDR-100, Campbell, UT, USA) with 40 cm probes installed in
the middle of each soil compartment. All the plants were kept well-
watered to pot holding capacity during the rst 44 days. Thereafter,
the plants were subjected to: (1) Full irrigation (FI): both soil com-
partments were watered to pot water holding capacity; (2) Partial
root-zone drying irrigation (PRD) treatments: half of the root sys-
tem was watered to pot water holding capacity while the other half
was allowed to dry to around 8% within about seven days before the
irrigation was switched between the two soil compartments; (3)
Decit irrigation (DI): the same amount of water used for the PRD
treatment was irrigated evenly to the two soil compartments. All
the pots were irrigated every second day until the end of the exper-
iment. The water used for irrigation was tap water with negligible
concentrations of nutrients.
68 C. Liu et al. / Agricultural Water Management 159 (2015) 6676
Both experiments had six treatments that consisted of a com-
bination of the two factors: P fertilization levels (P0 or P1) and
irrigation methods (FI, PRD or DI). Exp1 had 6 replicates and Exp2
had 4 replicates, both were arranged in randomized complete block
designs. Exp2 was conducted to further conrm the result of the
Exp1. The differences between Exp2 and Exp1 were that soil used
in Exp2 was sterilized to eliminate microbial inuence on water or
P uptake by the plants. The sterilization process of Exp2 released
some bioavailable nutrient into the soil, and therefore we reduced
the N and P fertilizer dose. Furthermore, there were three harvest
times in Exp1 to follow plant water and P utilization dynamics but
only one harvest time in Exp2, which was at the same potato devel-
opment stage as the second harvest of Exp1. The fertilizer used in
Exp1 was compound fertilizer while the fertilizer used in Exp2 was
pure chemical. The soil used in both experiments had relatively low
concentration of P to induce P deciency in the plants.
2.2. Sampling, measurements, calculations and statistical
analyses
The irrigation volumes (Wi ) were calculated at each irrigation
time based on TDR-measurements with two to four replicates of
each treatment using the formula:
W i (L) = Vs(L) ( p  a ) (1)
where: Vs is the soil volume in each pot,  p is the volumetric water
content (%) at pot water holding capacity and  a is the actual volu-
metric water content (%) as measured by TDR.
The applied water volume during the irrigation treatment was
the sum of the irrigation volumes (Wi ) at each irrigation time. Aver-
age soil volumetric water content () was calculated by averaging
the soil water content before and after irrigation.
In Exp1, the rst, second and nal harvests were done after 0, 27
and 52 (nal harvest) days irrigation treatments. Leaf water poten-
tial was measured one day before nal harvest with a pressure
chamber (Soil Moisture Equipment Corp., Santa Barbara, CA, USA)
on the second (from the top) fully expanded upper leaf. Shoots (leaf
and stem) and underground parts (root and tuber) of the plant, bulk
and rhizosphere soil were collected at each harvest. At nal harvest,
leaf area was measured with a leaf area meter (LI-3100C, Li-Cor Lin-
coln, NE, USA). The bulk soil was collected by mixing all the pot soil
(topsoil and subsoil) evenly. The rhizosphere soil was collected by
shaking roots vigorously to separate soil particles loosely adhering
to the fresh roots. Roots, tubers, stems and leafs of each harvest
were dried at 70 C for 72 h to constant weight and the dry biomass
was recorded. Total (dry) biomass is the sum weight of roots, tubers,
stems and leafs. Root to shoot ratio (RSR) was calculated as:
100 root dry biomass
RSR(%) = (2)
shoot dry biomass
Water use efciency (WUE g L1 ) was calculated as:
WUE(g L1 )
plant total biomass increment (g)
= (3)
applied water volume during irrigation treatment (L)
Plant samples were divided into shoots and underground parts,
and the dried samples were ground to a ne powder. Total P con-
centration was measured at each harvest time after ashing (450 C),
solubilizing in 13.9% hydrochloric acid and 21.7% nitric acid and
was analyzed using a spectrophotometer after the addition of
vanadate molybdate (Stufns, 1967). Total nitrogen (N) concen-
tration was measured at nal harvest with LECO CNS-1000 (LECO
Corp., St. Joseph, MI. USA) according to ISO 13,878. Plant P and N
uptake (mg plant1 ) was calculated as total P and N concentration
of plants (mg g1 ), respectively multiplied by plant biomass
(g plant1 ). Plant N:P ratio (NPR) was calculated as:
plant N uptake
NPR = (4)
plant P uptake
Agronomic P use efciency (APUE) was calculated as:
g biomass (Y1 Y0)
APUE( )= (5)
mg P fertilizer Pf
where: Y1 and Y0 are total biomass of P1 and P0 plants respectively;
Pf is the rate of fertilizer P applied in P1 treatments.
Water soluble P (WSP) in the soil was measured at each har-
vest time in pooled supernatants after two sequential extractions
of 1 g of fresh soil shaken with 50 mL of deionized water for 1 h at
20 C followed by centrifuging for 30 min at 10,733 g at 20 C and
was analyzed with a spectrophotometer at 890 nm after the addi-
tion of vanadate molybdate (Sissingh, 1971). Microbial biomass P
(MBP) was measured at each harvest time by measuring the resin P
difference with or without hexmo addition to soil. The resin P was
extracted with anion exchange membranes (AEM) where 2 g fresh
soil was suspended in 30 ml deionized water (Kouno et al., 1995).
Soil mineral N was only measured at nal harvest. The soil nitrate-N
(NO3 N) and ammonium-N (NH4 + -N) were extracted from 5 g fresh
soil with 50 ml 1 M KCl and then measured by an Autoanalyser III
(Bran + Luebbe, Germany). We assumed that P transport to root
surface was the sum of mass ow and diffusion. Percentage of P
transport to the root surface by mass ow (P transport massow ) and
diffusion (P transport diffusion ) was calculated according to Scheffer
and Schachtschabel (1979) as:
Pm
Ptransportmassow = 100 (6)
Phyp
Ptransportdiffusion = 100 Ptransportmassow (7)
where: Pm is the measured P concentrations which was taken as the
bulk soil WSP before start of irrigation treatments converted to mg
P/L soil solution according to Koopmans et al., (2010) and Blaauw
et al., (1988). Phyp is the calculated hypothetical P concentrations
in the soil solution if plant P uptake was achieved solely by mass
ow transport.
Pm = 0.00375 Pw 1.466 (8)
Phyp (mg/L)
plant P uptake
= (9)
water used by transpiration during irrigation treatment
where: Pw is the soil P concentration expressed by mg P2 O5 / L air
dried soil.
The soil mineral N balance (SNmin ) was calculated as the differ-
ence between the sum of soil mineral N (SNt1 ) and applied mineral
N (AN) at the onset of the experiment (t1 ) and the sum of bulk soil
mineral N (SNt2 ) and average plant biomass N (PN) at the time of
nal harvest (t2 ):
SNmin = (SNt1 + AN)(SNt2 + PN)(10)
In Exp2, the harvest was done 30 days after irrigation treatments
started. Plant and bulk soil samples were collected. Plant biomass,
WSP, total P concentration of plant, root to shoot ratio and P uptake
were measured and calculated in the same way as exp1. Soil acid
phosphomonoesterase (APM) activity was measured with a spec-
trophotometer at 420 nm (Tabatabai and Bremner, 1969) after 1 g
(dry weight) fresh soil had been incubated in a universal buffer at
37 C with p-nitrophenyl phosphate solution for one hour. Results
 C. Liu et al. / Agricultural Water Management 159 (2015) 6676 69

Table 1
Total biomass of plant (g plant1 ) as affected by P fertilization levels (P0: without P fertilizer, P1: with P fertilizer) after 0, 27 and 52 days (Exp1) and 30 days (Exp2) irrigation
treatments (FI: full irrigation, DI: decit irrigation and PRD: partial root zone drying irrigation). There was a signicant interaction of P fertilization levels and irrigation
methods (P < 0.05) at harvest after 52 days.

Irrigation 0 day (Exp1) 27 day (Exp1) 52 day (Exp1) 30 day (Exp2)

method
P0 P1 P0 P1 P0 P1 P0 P1

FI 15.7 B 25.8 A 36.0 B 66.9 A 64.9aB 150.3aA 64.3aB 71.2aA

DI 29.9 B 58.4 A 64.1aB 112.0bA 55.9bB 63.9bA
PRD 34.3 B 65.1 A 62.1aB 115.8bA 58.9b 62.7b

Values are means (Exp1: n = 6, Exp2: n = 4).

Signicant differences are indicated by letters: Different small letters within a column indicate signicant difference due to irrigation method at P < 0.05 within same P level.
Different capital letters within a row within same harvest time indicate signicant difference due to P fertilization level at P < 0.05 within same irrigation method.

were calculated as g p-nitrophenol released per gram dry soil per
hour.
The statistical analyses were conducted using the MIXED pro-
cedure of SAS (9.3). The xed effects were irrigation treatments, P
fertilization levels and irrigation treatments P fertilization levels.
Replicate was included as a random effect. The correlation analysis
was done by the CORR procedure of SAS (9.3).
3. Results
3.1. Plant growth, soil water dynamics and water use efciency
under different irrigation regimes
The application of phosphorus (P) fertilizer increased potato
growth in both experiments, but to a larger extent in Exp1 than
Exp2 (Table 1), possibly due to the higher P-availability in the P0
treatment of Exp2 caused by the soil sterilization. In both exper-
iments under P1, the total plant biomass under decit irrigation
(DI) and partial root zone drying irrigation (PRD) treatments was
similar, and was about 24% lower than that of full irrigation (FI).
However, at P0 in Exp1, no difference of plant biomass was found
for the three irrigation regimes, giving rise to an interactive effect
between P and irrigation at nal harvest, while in Exp2, the total
biomass of DI and PRD was lower than FI both at P0 and P1.
The application of P fertilizer also increased the root biomass in
both experiments (Fig. 2). Further, the root biomass was affected by
irrigation treatments but differently between the two experiments
as root biomass in Exp1 was lower under DI and PRD compared to
FI, while root biomass in Exp2 was higher under DI and FI compared
to PRD. The effects were generally larger under P1 than P0 giving

Fig. 2. A: Root biomass (g plant1 ) and root to shoot ratio (%) as affected by P fertilization levels (P0: without P fertilizer, P1: with P fertilizer) after 0, 27 and 52 days (Exp1)
and 30 days (Exp2) irrigation treatments (FI: full irrigation, DI: decit irrigation and PRD: partial root zone drying irrigation). B: leaf water potential (MPa) and leaf area (cm2
plant1 ) as affected by P fertilization levels (P0: without P fertilizer, P1: with P fertilizer) after 52 days (Exp1) irrigation treatments (FI: full irrigation, DI: decit irrigation and
PRD: partial root zone drying irrigation). Error bars indicate SE (n = 6 in Exp1, n = 4 in Exp2). Signicant differences are indicated by letters: grouped columns without letters
in common are signicantly different between irrigation treatments at P < 0.05. Symbol [Special symbol replaced] inside top of column indicates signicantly higher value
of the variable due to P fertilization level (P < 0.05) within same irrigation method and harvest time. There were signicant interactions of P fertilization levels and irrigation
methods on root biomass at 30 days harvest and on leaf area at 52 days harvest.
70 C. Liu et al. / Agricultural Water Management 159 (2015) 6676

Fig. 3. Average soil volumetric water content under three irrigation treatments (FI: full irrigation, DI: decit irrigation and PRD: partial root zone drying irrigation) during
52 days (Exp1) and 30 days (Exp2) irrigation treatments. In PRD, two soil compartments: PRD-L and PRD-R were recorded. Values are means (n = 24).

rise to a signicant interaction effect between P and irrigation with Table 2

Water use efciency (WUE g/L, total plant biomass increment/plant water use) as
respect to root biomass in Exp2. The application of P fertilizer also
affected by P fertilization levels (P0: without P fertilizer, P1: with P fertilizer) after
increased potato leaf expansion but generally P deciency induced 27 and 52 days (Exp1) irrigation treatments (FI: full irrigation, DI: decit irrigation
higher root to shoot ratio (Fig. 2). Water stress induced by DI and and PRD: partial root zone drying irrigation).
PRD did not induce higher root to shoot ratio in Exp1. At P1, PRD
Irrigation 27 day 52 day
caused a lower leaf water potential than DI and FI. method
P0 P1 P0 P1
The dynamic soil water content () in the 040 cm soil prole
in the two experiments (Exp1 and Exp2) is shown in Fig. 3. In Exp1 FI 6.51 5.13b 5.41b 6.17b
in P1pots, the mean  of FI and DI was about 21% and 15%, respec- DI 5.08 6.54ab 6.89a 6.83ab
PRD 6.67 7.89a 6.61a 7.12a
tively, while for PRD,  decreased progressively from 19% to 9% in
the dry compartments within about 7 days and with a mean  of Values are means (Exp1: n = 6, Exp2: n = 4).
Signicant differences are indicated by letters: Different letters within a column
16% over both the dry and wet side. FI was irrigated with 8 L and
indicate signicant difference due to irrigation method at P < 0.05 within same P
20.2 L water and DI and PRD was irrigated with 5 L and 12.6 L water level.
when harvested at 27 and 52 day, respectively. In P0 pots, the mean
of FI and DI was about 23% and 17%, respectively, while for PRD,
 decreased steadily over time from around 22% to 12% in the dry  decreased steadily over time from around 18% to 9% in the dry
compartments within about 7 days and with a mean  of 19% over compartments within about 7 days and with a mean  of 15 % over
both the dry and wet side. FI was irrigated with 3.1 L and 9.1 L water both the dry and wet side. FI was irrigated with 10 L and 8.8 L water
and DI and PRD was irrigated with 2.8 L and 7 L water when har- and DI and PRD was irrigated with 6.8 L and 6.7 L water for P1 and
vested at 27 and 52 day, respectively. As bare soil evaporation was P0, respectively harvested at 30 day.
measured to have a negligible value compared with total water In Exp1, when harvested at 52 day at P1, PRD and DI plants had
usage, we assumed that all the irrigation water was used for plant saved 37% water and improved WUE with respect to tuber biomass
transpiration. The application of P fertilizer had no effect on water by 29% under PRD compared with FI but at the expense of 23%
use efciency (WUE). Under P0, WUE was similar under PRD and and 25% reduction in total biomass and 16% and 21% reduction
DI treatments which were signicant larger than FI at nal harvest in tuber biomass. At P0, PRD and DI plants had saved 23% water
(Table 2). But under P1, only PRD had signicantly larger WUE than and improved WUE with respect to tuber yield by ca. 35% as total
FI treatments. biomass and tuber biomass were not reduced compared to FI.
In Exp2 in P1 pots, the mean  of FI and DI was about 21% and
16%, respectively, while for PRD,  decreased steadily from around 3.2. Plant P acquisition and P dynamics in the root zone
19% to 11% in the dry compartments within about 7 days and with a
mean  of 17% over both the dry and wet side. In P0 pots, the mean The application of P fertilizer increased plant P concentration
 of FI and DI was about 22% and 15%, respectively, while for PRD, by about 20% but almost tripled the P uptake at nal harvest
 C. Liu et al. / Agricultural Water Management 159 (2015) 6676 71

Fig. 4. Potato P concentration (mg g1 ) and P uptake (mg plant1 ) as affected by P fertilization levels (P0: without P fertilizer, P1: with P fertilizer) after 0, 27 and 52 days
(Exp1) and 30 days (Exp2) irrigation treatments (FI: full irrigation, DI: decit irrigation and PRD: partial root zone drying irrigation). Error bars indicate SE (n = 6 in Exp1, n = 4
in Exp2). Signicant differences are indicated by letters: grouped columns without letters in common are signicantly different between irrigation treatments at P < 0.05.
Symbol [] inside top of column indicates signicantly higher value of the variable due to P fertilization level (P < 0.05) within same irrigation method and harvest time.
There were signicant interactions of P fertilization levels and irrigation methods on shoot P concentration, shoot P uptake and total P uptake at 52 days harvest and on shoot
P uptake at 30 days harvest.

(Fig. 4). The sequential harvests showed that plant P concentration
decreased during plant growth. Under P1, P concentration of shoots
initially was higher under FI than DI and PRD, later became almost
equal to DI and PRD, and nally decreased and became lower than
DI and PRD when harvested at 52 day. At P0 levels in Exp1, shoot
P concentration of PRD was higher than FI and DI when harvested
at 52 day, while PRD had similar biomass production as DI. In both
experiments at P1 levels, DI and PRD decreased P uptake by about
22% compared with FI at nal harvest, and P uptake of DI and PRD
was similar. All the three irrigation treatments had similar P uptake
at P0 levels when harvested at 52 day in Exp1, giving rise to a signif-
icant interaction between P and irrigation treatments. In contrast, P
uptake in Exp2 at P0 and P1 levels showed the same pattern for the
three irrigation treatments with DI and PRD being lower than FI.
P concentration and plant uptake were higher at P0 levels in Exp2
than in Exp1 as mentioned earlier possibly due to the soil steriliza-
tion in Exp2. The P uptake positively correlated to plant root dry
biomass (R2 = 0.6, P < 0.001) (Fig. 5), while the P concentration of
plant negatively correlated to total biomass at high P level (R2 = 0.8,
P < 0.001) (Fig. 5).
The application of P fertilizer increased water soluble P (WSP)
and microbial biomass P (MBP) but decreased activity of acid phos-
phomonoesterase (APM) (Fig. 6). WSP in rhizosphere soil decreased
along with potato growth at P1 levels, while it was much lower and
almost constant at the P0 level. WSP in bulk soil showed similar
dynamics as in rhizosphere soil but with smaller values (data not
shown). When harvested at 27 (Exp1) and 30 day (Exp2), WSP of
DI was higher than PRD in both rhizosphere and bulk soil. When
harvested at 52 day, WSP of FI was lower than DI and PRD (resin
P had similar dynamic as WSP, but the value was 4-5 times larger;
data not shown). MBP also had the tendency to increase along with
the growth of potato (Fig. 6; Exp1) and MBP in rhizosphere of DI
was larger than FI and PRD. APM concentration of PRD was higher
than DI and also tended to be higher than FI at both P levels (Fig. 6;
Exp2).
Agronomic P use efciency (APUE g biomass/g P applied per
plant) was signicantly inuenced by irrigation treatments at nal
harvest in Exp1. The APUE (g biomass/g P applied per plant) of FI, DI
and PRD were 107, 60 and 67, respectively. The APUE of FI treatment
was much larger than DI and PRD, while APUE of PRD was slightly
higher than DI. Generally, soil P transport to the root surface was
dominated by diffusion, which accounted for more than 95% of the
total P uptake (Fig. 7). The application of P fertilizer decreased the
proportion of P transported to the root surface by diffusion. The
proportion of P transported to the root surface by diffusion was
similar under DI and PRD and higher than under FI (Fig 7).
3.3. Effect of irrigation on plant N uptake, plant N:P ratio and soil
mineral N balance
The application of P fertilizer strongly increased potato N uptake
but decreased plant N:P ratio (Table 3). Shoot and total N uptake
under PRD was higher than under DI at P1 and also tended to be
higher than DI at P0. While shoot and total N uptake were highest
under FI at P1, this was strikingly different at P0 levels where PRD
had the highest N uptake, and therefore there was a signicant
interaction between P levels and irrigation treatment. Shoot and
total N:P ratio of PRD tended to be higher than in DI and FI, and this
72 C. Liu et al. / Agricultural Water Management 159 (2015) 6676

Fig. 5. (A) Liner regression between shoot P concentrations: total biomass at P fertilization levels of P1 (with P fertilizer) under three irrigation treatments (FI: full irrigation,
DI: decit irrigation and PRD: partial root zone drying irrigation) and three harvest times 0, 27 and 52 days (Exp1). (B) Liner regression between P uptake: root biomass at P
fertilization levels (P0: without P fertilizer, P1: with P fertilizer) under three irrigation treatments (FI: full irrigation, DI: decit irrigation and PRD: partial root zone drying
irrigation) and three harvest times 0, 27 and 52 days (Exp1) and 30 days (Exp2).

effect was larger at P1 than P0 levels and had a trend to increase
with declining soil water content at both high and low P levels
(curve not shown).
It was clear that P fertilizer decreased mineral N left in soil
(Table 4). Soil NO3 -N concentration in PRD and DI was similar and
was higher than in FI in both rhizosphere and bulk soil. For the
P0 levels, higher N uptake of PRD compared to FI was observed.
The NH4 -N concentrations of rhizosphere and bulk soil resembled
those of NO3 -N but DI was higher than PRD (P = 0.059). At the begin-
ning of the experiment, NO3 -N and NH4 -N were 145 mg kg1 and
132 mg kg1 soil, respectively. After harvest, NO3 -N had decreased
to 235 mg kg1 and NH4 -N had decreased to 03 mg kg1 .
The application of P fertilizer also strongly decreased the soil
mineral N balance (SNmin ) (Table 4). SNmin was highest under

Fig. 6. Exp1: Rhizosphere soil water soluble P (WSP, mg kg1 ) and microbial biomass P (MBP, mg kg1 R = 82.091.2% percentage recovery of added P fertilizer) as affected by
P fertilization levels (P0: without P fertilizer, P1: with P fertilizer) after 0, 27 and 52 days (Exp1) irrigation treatments (FI: full irrigation, DI: decit irrigation and PRD: partial
root zone drying irrigation). Error bars indicate SE (n = 6). Exp2: Bulk soil water soluble P (WSP) and acid phosphomonoesterases (APM) activity as affected by P fertilization
levels (P0, P1), irrigation treatments (FI, DI and PRD) after 30 days (Exp2). Error bars indicate S.E. (n = 4). Signicant differences are indicated by letters: grouped columns
without letters in common are signicantly different between irrigation treatments at P < 0.05. Symbol [Special symbol replaced] inside top of column indicates signicantly
higher value of the variable due to P fertilization level (P < 0.05) within same irrigation method and harvest time. There were signicant interactions of P fertilization levels
and irrigation methods on rhizosphere soil WSP and MBP at 27 days harvest.
 C. Liu et al. / Agricultural Water Management 159 (2015) 6676 73

we found PRD and decit irrigation (DI) had an equally reduced P

Fig. 7. Percentage of P transport to root surface by diffusion in total P transport (%) as
affected by P fertilization levels (P0: without P fertilizer, P1: with P fertilizer) after 27
and 52 days (Exp1) irrigation treatments (FI: full irrigation, DI: decit irrigation and
PRD: partial root zone drying irrigation). Error bars indicate SE (n = 6). Signicant
differences are indicated by letters: grouped columns without letters in common
are signicantly different between irrigation treatments at P < 0.05. Symbol []
inside top of column indicates signicantly higher value of the variable due to P
fertilization level (P < 0.05) within same irrigation method and harvest time. There
were signicant interactions of P fertilization levels and irrigation methods at 27
and 52 days harvests.
FI at P0 levels where it amounted to 46% of the N fertilizer added at
the start of the experiment, but at P1 levels it was lowest under FI
and thus an interactive effect of P and irrigation was found. At both
P levels however, SNmin was larger under DI than under PRD.
4. Discussion
It was hypothesized that partial root zone drying irrigation
(PRD) can improve nitrogen (N) uptake, and the improved N uptake
might be accompanied by an improved phosphorus (P) uptake.
The result showed that compared to DI, PRD improved N uptake.
However, we did not nd improved P uptake under PRD, instead
uptake at P1 level compared with full irrigation (FI), while DI, PRD
and FI had similar total P uptake at P0 level. Nevertheless, consistent
with previous studies (Liu et al., 2006; Shahnazari et al., 2008; Wang
et al., 2010a,b, 2013; Sun et al., 2013), similar decreased potato
plant biomass and leaf expansion, improved water use efciency
(WUE) of PRD and DI compared with FI and increased N uptake of
PRD compared with DI were found in this study.
The soil used in these two pot experiments was a low P soil;
indeed the P concentration in P0 soil was so low and constituted
such strong limitation on plant growth that irrigation methods had
negligible inuence on the P-uptake of P0 plants. The result of the
second experiment conrmed the results of the rst experiment,
except the soil sterilization released bioavailable P as indicated by
increased water soluble P (WSP) in soil and improved P concen-
tration and uptake in plants, so P0 plants in Exp2 did not show
similar severe lack-of-P symptoms as in Exp1, but still differences in
P-uptake were small and non-signicant between irrigation treat-
ments.
Plant P uptake and P use efciency may be improved by: (i) roots
exploring a larger soil volume, (ii) easing transport of P from bulk
soil to root surface and (iii) transforming non-available P forms into
plant-available P (Rengel, 2008). In the current study, the P fertilizer
was applied only to the top 2022 cm soil in both experiments. As
the P easily binds to soil particles, downward movement of P under
these experimental conditions would be small, and the fertilizer
P would be concentrated in the top soil layer; the same situation
as usually found under eld conditions. The factors that affect P
uptake i.e. root development and function, P diffusion in soil and
plant available P, are discussed in the following paragraphs.
Root architecture plays an important role in P acquisition
(Gahoonia, 1994; Lynch, 1995) because it determines the size of
the explored volume of soil. The repressed primary root growth
in response to P deciency conditions is based on the presence of
NO3 (Medici et al., 2015), clearly indicating an integrated response
of root architecture formation to nitrate and phosphate. Consistent
with Hodge, (2004), an enhanced root growth due to P fertiliza-
tion was found in the current study, and the P uptake exhibited a

Table 3
Plant N uptake and plant N: P ratio as affected by P fertilization levels (P0: without P fertilizer, P1: with P fertilizer) after 52 days (Exp1) irrigation treatments (FI: full irrigation,
DI: decit irrigation and PRD: partial root zone drying irrigation). There was a signicant interaction of P fertilization levels and irrigation methods (P < 0.05) on shoot and
total N uptake.

Irrigation Plant N uptake (mg/pot) Plant N:P ratio

method
Shoot Total plant Shoot Total plant

P0 P1 P0 P1 P0 P1 P0 P1

FI 571bB 1771aA 1483bB 3125aA 26.1b 23.8b 18.5 A 13.7 B

DI 632abB 1217cA 1640abB 2654cA 32.0abA 21.6bB 20.3 A 14.8 B
PRD 844aB 1500bA 1706aB 2872bA 34.5a 29.3a 20.1 A 16.2 B

Values are means (for plant N uptake: n = 3, for plant N: P ratio: n = 6).
Signicant differences are indicated by letters: Different small letters within a column indicate signicant difference due to irrigation method at P < 0.05 within same P level.
Different capital letters within a row within same harvest time indicate signicant difference due to P fertilization level at P < 0.05 within same irrigation method.

Table 4
Soil mineral N (NO3 -N, NH4 -N) left in soil and mineral N balance (SNmin , mg/pot) as affected by P fertilization levels (P0: without P fertilizer, P1: with P fertilizer) after 52
days (Exp1) irrigation treatments (FI: full irrigation, DI: decit irrigation and PRD: partial root zone drying irrigation).

Irrigation Soil NO3 -N (mg/kg soil) Soil NH4 -N (mg/kg soil) Soil SNmin (mg/pot)
method
Rhizosphere Bulk Rhizosphere Bulk

P0 P1 P0 P1 P0 P1 P0 P1 P0 P1

FI 22.2bA 6.4bB 28.0bA 2.5bB 0.88b 0.40b 0.63 A 0.25 B 1379aA 27cB
DI 34.7aA 18.9aB 31.6aA 15.2aB 2.47a 1.68a 0.82A 0.45 B 1181bA 355aB
PRD 33.3aA 19.7aB 31.6aA 13.8aB 1.85a 1.33a 0.52 0.40 1118cA 152bB

Values are means (n = 6).

Signicant differences are indicated by letters: Different small letters within a column indicate signicant difference due to irrigation method at P < 0.05 within same P level.
Different capital letters within a row within same harvest time indicate signicant difference due to P fertilization level at P < 0.05 within same irrigation method.
74 C. Liu et al. / Agricultural Water Management 159 (2015) 6676
signicantly positive linear relationship with root biomass.
Although no root mass difference was found in previous studies in
potato irrigated with PRD, DI and FI (Ahmadi et al., 2011). FI had the
highest root mass and PRD restricted root growth compared with
DI (only in Exp2). An increased root to shoot ratio was found under
P0, which is a typical response to P decient condition (Wissuwa
et al., 2005), as preferential assimilate distribution to the roots
potentially may increase the plant P uptake. The pots used in the
experiment were 16 cm in diameter, which could have affected the
measured responses with respect to plant biomass, root biomass
etc. Compared with plants grown in the eld, the potted plants had
less soil volume for root growth. Also, the arrangement of plants in
the greenhouse left more space for top growth, which may affect
plant shape, light interception and transpiration rate. As a result,
the total biomass, root biomass, N uptake, P uptake and tuber pro-
duction might be different from plants grown in eld. The effects
on root distribution and root length density might have affected P
uptake (Miguel et al., 2015), and therefore further experiments are
needed to verify their role on P uptake. Diffusion, the main pro-
cess making P available to roots, supplies P from the bulk soil to
the root surface (Comerford, 1998). Current results showed that,
diffusion transferred more than 95% of P from soil to roots. There-
fore, no matter the kind of irrigation treatment used, diffusion was
the main way of P transport to the root surface, which implied
that the percentage of P transport to the root surface by diffusion
was less affected by irrigation pattern than by soil water content.
Plant growth is also an important factor affecting P uptake (White
et al., 2005b) as enhanced root growth will increase the uptake
by increasing the explored soil volume and cross sectional area of
diffusion. So any other environmental factors that improve root
growth may also be accompanied with a higher P uptake.
Irrigation using the PRD method is accompanied by drying and
rewetting cycles (D/RW) of the soil prole. D/RW causes microbial
death and cell lysis (Turner et al., 2003), followed by an increase
in soil respiration rate indicating enhanced microbial activity and
ushes of mineral N and P into soil solution (Birch 1958; Xiang et al.,
2008; Butterly et al., 2009). However, other results have indicated
that soil available phosphate is not affected by wetting a dry soil
(Cui and Caldwell, 1997). The soil microbial biomass plays a very
important role in soil fertility both as a source and sink for mineral
nutrients (Kouno et al., 1995). In the present study, D/RW of PRD
caused a reduction in the size of microbial biomass compared to DI,
measured as biomass P which is consistent with previous studies
(Wu and Brookes, 2005; Gordon et al., 2008). The choice of dry-
ing to 8% (Vol) was based on the soil water characteristics curve
and results from the article of Liu et al., (2008) suggesting that the
dry side should be allowed to dry to a water potential of around
80 kPa to induce relevant ABA production in potatoes. In current
experiments, except from reduced size of microbial biomass, there
was additional evidence of severe stress caused by D/RW of PRD as
indicated by lower leaf area, total biomass, root biomass and leaf
water potential, all of which implied that allowing the water con-
tent to decrease to 8% (Vol) for the dry side of PRD before switching
between the sides was too extreme to maintain growth. However,
no ush of plant available P under PRD was found in this study com-
pared with DI. When given same amount of P fertilizer, WSP, which
is plant available, decreased along with plant P uptake. As a result,
it was inappropriate to compare plant available P in case the plant
P uptake was different. Higher P availability (soil WSP and resin
P) was found under DI instead of PRD with high P levels at 27 day
(Exp1) and 30 day (Exp2) harvest, which may indicate a xation of
available P by D/RW (Fig. 6).
Organic P (Po) accounts for 5070% of the total P in soils (Borie
and Rubio, 2003). Phosphatase catalyzing the hydrolysis of Po is
important to soil P availability. Soil moisture affects the soil microbe
and root activity and is an important factor affecting soil enzyme
activity. A 10% reduction of soil moisture may decrease acid phos-
phatase activity by 3040% (Sardans and Penuelas, 2005). Acid
phosphomonoesterase activity was only measured in Exp2 with
sterile soil in this study, so it was assumed that they were secreted
by plant roots. The results showed that acid phosphatase activity
was decreased by DI treatments compared to FI (Fig. 6), but PRD
treatment had root acid phosphatase activity similar to FI, which
were higher than DI. This nding differed from results of previ-
ous work (Li et al., 2010) where PRD decreased the activity of acid
phosphatase.
In a review of 40 fertilization studies it was concluded that, N:P
ratios in plants >16 are indicative of P limitation, while N: P < 14
indicate N limitation. Ratios of N:P between 14 and 16 indicate
either N or P or possibly N and P together are limiting plant growth
(Koerselman and Meuleman, 1996). The N:P ratios reect the rel-
ative availability of N and P to plants and may indicate the degree
of N or P deciency experienced by a plant (Gusewell et al., 2003).
According to this standard, as can been seen from Table 3, all the
irrigation treatments at the low P levels suffered a severe P de-
ciency, while treatments at the high P levels under FI suffered slight
N deciency, and PRD suffered slight P deciency. Shoot N:P ratio
also indicated that PRD treatments went through a more severe
P deciency than DI and FI and this deciency might have trig-
gered more root phosphatase secretion in PRD than in DI. The higher
phosphatase activity of PRD might have resulted in a higher min-
eralization of Po under PRD than DI. However this Po contribution
to plant total P uptake was not enough to result in a difference
of P uptake under DI and PRD, since WSP tended to be higher in
DI and ultimately similar P uptake was achieved under DI and
PRD. The trend that shoot N:P ratio increased with declining soil
water content was in agreement with Cernusak et al. (2010) and
was explained by differential changes in the mobility of nitrate vs
phosphate in response to declining soil water content.
In the present study, we found improved N uptake of PRD both
at low and high P levels compared to DI. The soil mineral N balance
(SNmin ) as dened here measures the decrease in soil mineral N
from planting to harvest other than explained by plant N uptake.
As SNmin was positive, there was a reduction of mineral N rather
than soil N mineralization as found in the study by Sun et al. (2013).
The possible reasons why SNmin was positive in PRD, DI and FI
were either denitrication, ammonia volatilization or immobiliza-
tion through microbial conversion of N to organic forms (Cameron
et al., 2013). Of these, immobilization of N to forms which are tem-
porarily unavailable to plants was the most probable cause for the
different SNmin under DI and PRD, since denitrication was likely
low due to the soil water content being kept at or below 70% eld
capacity as well as the low soil pH (below 7) was not favoring
NH3 volatilization. There are substantial indirect and direct evi-
dence supporting that the competition exists between plants and
microorganisms (Kaye and Hart, 1997) and the applied inorganic N
can be rapidly taken up by the soil microbes (Nannipieri et al., 1990).
Up to 50% immobilization of fertilizer N has also been found in other
pot and eld studies (Blankenau et al., 2000) and in the present
experiments seemed to be favored by the high concentration of
mineral N in the soil under P0 conditions where plant uptake was
limited by P deciency. Also the use of ammonium nitrate as fer-
tilizer may have favored immobilization processes as ammonium
seems to be the preferred substrate for bacterial immobilization
(Wickramasinghe et al., 1985; Blankenau et al., 2002). The lower
SNmin found under PRD than under DI in P1 may also suggest less
N immobilization in PRD and this corresponded to the higher plant
N uptake under PRD. The tendency of larger root biomass especially
under P1 which can provide carbon for microbial metabolism also
corresponded to the tendency of higher MBP in DI than PRD pots.
The estimation of mineral N immobilization under DI and PRD at
P1 tted the value of SNmin by the calculation of soil microbial
 C. Liu et al. / Agricultural Water Management 159 (2015) 6676
biomass N based on average soil microbial biomass N:P ratios of
7:1 in a meta-analysis at the global scale by Cleveland and Liptzin
(2007). The results thus implied that higher microbial biomass in DI
enhanced mineral N immobilization, and caused stronger compe-
tition with plants for limited quantities of mineral N as compared
to PRD. This nding was similar to the previous studies that PRD
plants have higher N uptake (Wang et al., 2010b, 2013), but at the
same time, differs in its conclusion as enhanced N uptake in pre-
vious studies was attributed to enhanced organic N mineralization
under PRD.
To sum up, DI and PRD triggered different physiological and
biochemical process in soil/ rhizosphere-plant continuum: The
D/RW under PRD introduced more severe stress than under DI and
resulted in reduced leaf water potential, root biomass and MBP.
D/RW in PRD might also lead to the xation of available P mea-
sured as lower WSP and resin P than DI. But an acclimation to P
deciency was triggered by higher plant N: P ratio in PRD indicated
by increased root secretion of acid phosphatase. All these plant and
microbial activity acclimations to scarcity of P and soil moisture
eventually induced a similar potato P uptake by PRD and DI and
higher N uptake by PRD than DI. The cause of higher N uptake of
the PRD plants might be attributed to the restricted soil microbial
activity and reduced microbial competition for mineral N induced
by the drying and rewetting cycles. When plants were subjected
to limited water and P at the same time as in the current study, P
was the most limiting factor affecting plant growth, and the P de-
cient condition induced high soil N immobilization and low plant
utilization of water and N fertilizer.
In conclusion, when same amount of water was used, PRD was
superior to DI in terms of N uptake, but was not distinctively differ-
ent from DI with respect of plant biomass production and P uptake.
Challenges remain as to maintaining crop yield and P uptake under
the reduced irrigation regimes used in contemporary water saving
irrigation strategies. Further studies on how water saving irrigation
strategies affects crop yield, root growth and P uptake under eld
conditions are required and remedial actions should be explored to
mitigate these negative effects during water management.
Acknowledgements
We thank Finn Henning Christensen, Mette Nielsen, Jens B.
Kjeldsen, Karin Dyrberg and Stig T. Rasmussen for their technical
assistance and Zhenjiang Zhou for helping to irrigate and harvest
plants. The Graduate School of Science and Technology (GSST),
Aarhus University contributed with funding for the project.
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