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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 half of the root zone is irrigated while the second half is exposed to
soil drying to a predetermined level before switching the irrigation
The increasing worldwide competition for fresh water resources to this second half. To maximize potato yield, not only sufcient P
requires better management of irrigation to improve water use ef- but also sufcient water need to be available (White et al., 2005a).
ciency (WUE). Decit irrigation (DI) and alternate partial root-zone Since DI and PRD reduce the soil moisture content, the tortuosity
drying irrigation (PRD) are two water saving irrigation regimes, of the diffusion pathway for nutrients increase (Gahoonia et al.,
which can increase WUE of potatoes (Wang et al., 2009) even with- 1994) and the transport of P from soil to root is reduced (McBeath
out reducing yield (Shahnazari et al., 2007). Under DI, the crop et al., 2012). Compared with DI, PRD increases the nitrogen (N)
receives irrigation water amounts slightly less than actual evap- availability by allegedly inducing greater microbial activity and
otranspiration but the resulting mild stress has minimal effects on mineralization of organic N, which resulted in improved N uptake
yield (English and Raja, 1996). PRD is a special form of DI where only 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,
Abbreviations: P, phosphorus; N, nitrogen; FI, full irrigation; DI, decit irrigation;
2007). Potato has a shallow root system (Vos and Haverkort, 2007),
PRD, partial root-zone drying irrigation; WUE, water use efciency; WSP, water sol- is generally drought sensitive (Yuan et al., 2003), and has a high
uble phosphorus; P1, treatments with phosphorus fertilizer; P0, treatments without phosphorus (P) requirement (Westermann, 2005). P is an essential
phosphorus fertilizer; MBP, microbial biomass P; SNmin , soil mineral N balance; element for all plants and rock phosphate from which P fertiliz-
APM, soil acid phosphomonoesterase activity.
Corresponding author at: Aarhus University, Blichers All 20, 8830 Tjele,
ers are derived is a vital non-renewable resource, which may be
Denmark. Tel.: +45 87154761.
depleted in a relatively short time span (Elser and Bennett, 2011).
E-mail address: caixialiu21@hotmail.com (C. Liu). 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 67

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., Fig. 1. Soil water retention curve for the experimental soil sampled from Ap-horizon
2001) and tomato (Wang et al., 2012) have been conducted, a thor- 030 cm at Foulumgaard.

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 removed from the tuber. In Exp2, potato tubers were surface
effect of FI, DI and PRD irrigation on P and N uptake in situations sterilized with 0.5% (v/v) H2 O2 for 1 min and rinsed thoroughly
with or without P fertilization. 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
2. Materials and methods initially placed outside under a transparent roof for 20 days and
then moved into the greenhouse. The conditions of the green-
2.1. Experimental materials and design 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
Two pot experiments were carried out in a climate-controlled photosynthetic active radiation (PAR) supplied by sunlight plus
greenhouse at Aarhus University Research Centre Foulum, metal-halide lamps. Pots were randomized on two tables in the
Denmark using cylindrical pots with a volume of 10 L (16 cm outer greenhouse.
diameter and 50 cm deep), divided into two vertical compartments In Exp1, two P fertilization levels were created: P1 in which the
by a diametric plastic sheet. A 5 cm wide and 10 cm high piece was 20 cm topsoil layer of each pot was supplemented with compound
cut and removed from the middle of the sheet at the top where the fertilizer consisting of 3.1 g N (47.73% NH4 -N, 52.27% NO3 -N), 2.57 g
seed tuber was going to be planted. Thus water exchange between K, 0.63 g S and 0.8 g P (CaHPO4 2H2 O); the control (P0) had the
the two compartments was prevented. The bottom of the pots was same amounts of N, K, S but no P. The calculation of fertilizer
covered by 1.5 mm nylon mesh. rates per plant was based on an assumed eld N application rate of
For the rst experiment (Exp1), the soil was collected from the 150 kg N ha1 and P rate of 37 kg P ha1 and further assuming that
experimental farm of Foulumgaard also at Research Centre Foulum, potatoes are planted at 40,000 plants ha1 (Haverkort, 1982). The
and was sandy loam, with 68% sand, 24.3% silt and 7.7% clay, total packed soil had a water content of 24.4% (v/v) at pot water holding
C 16.3 g Kg1 , total N 1.5 g Kg1 , Olsen P 22 mg kg1 , water solu- capacity measured in situ after 2 days of drainage of the pots placed
ble phosphorus (P) (WSP) 6.3 mg kg1 , mineral N:(NH4 + and NO3 ) on a moist, naturally drained outdoor soil surface and 5.18% (v/v) at
7.4 mg kg1 , pH 5.4. The eld capacity and wilting point of the soil permanent wilting point as measured in a pressure plate apparatus
at pF = 2.0 and pF = 4.2 were 25.3% (Vol.) and 6.7% (Vol.), respec- (Fig. 1). In Exp2, P1 was supplemented with 2.6 g N, 2.8 g K, 1.8 g S
tively. The water retention curve for the soil is shown in Fig. 1. The and 0.6 g P mixed into the 22 cm topsoil, while the control (P0) had
soil was dried under a roof to a water content of ca. 12% (v/v) and no P but otherwise the same dressing. Fertilizers used were pure
sieved through a 1 cm mesh sized sieve. Each pot was lled to a chemicals: NH4 NO3 , KH2 PO4 , K2 SO4 , MgSO4 7H2 O. The packed soil
dry bulk density of 1.3 g cm3 corresponding to a total of 11.17 kg had a water content of 23.8% (v/v) at pot water holding capacity.
dry soil. The soil was divided into a subsoil layer with a depth of Soil water content was monitored by a time domain reectome-
24 cm that was unfertilized and a topsoil layer with depth of 20 cm ter (TDR-100, Campbell, UT, USA) with 40 cm probes installed in
that was mixed with fertilizer before packing. In the second exper- the middle of each soil compartment. All the plants were kept well-
iment (Exp2), the sandy loam soil was collected from the same site watered to pot holding capacity during the rst 44 days. Thereafter,
as Exp1 with Olsen P 22 mg kg1 , WSP 8.7 mg kg1 , pH 6.2. The soil the plants were subjected to: (1) Full irrigation (FI): both soil com-
was air dried and sieved (1 cm mesh) and heated in an oven two partments were watered to pot water holding capacity; (2) Partial
times for 24 h at 85 C with an interval of 48 h at room tempera- root-zone drying irrigation (PRD) treatments: half of the root sys-
ture to eliminate living microorganisms. Sterilized soil had Olsen tem was watered to pot water holding capacity while the other half
P 26 mg kg1 , WSP 12 mg kg1 , pH 6.0. Pots were pre-sterilized by was allowed to dry to around 8% within about seven days before the
washing with 2% (w/v) NaClO and rinsed with tap water. Each pot irrigation was switched between the two soil compartments; (3)
was lled with 10.84 kg dry soil to a dry bulk density of 1.3 g cm3 Decit irrigation (DI): the same amount of water used for the PRD
with topsoil and subsoil layer of 22 cm each. treatment was irrigated evenly to the two soil compartments. All
Seed potatoes (Solanum tuberosum L. cv. Folva, size: 5580 g) the pots were irrigated every second day until the end of the exper-
were pre-germinated. When sprouts emerged, one seed potato iment. The water used for irrigation was tap water with negligible
was planted in the middle of each pot with all except one sprout concentrations of nutrients.
68 C. Liu et al. / Agricultural Water Management 159 (2015) 6676

Both experiments had six treatments that consisted of a com- of plants (mg g1 ), respectively multiplied by plant biomass
bination of the two factors: P fertilization levels (P0 or P1) and (g plant1 ). Plant N:P ratio (NPR) was calculated as:
irrigation methods (FI, PRD or DI). Exp1 had 6 replicates and Exp2
plant N uptake
had 4 replicates, both were arranged in randomized complete block NPR = (4)
plant P uptake
designs. Exp2 was conducted to further conrm the result of the
Exp1. The differences between Exp2 and Exp1 were that soil used Agronomic P use efciency (APUE) was calculated as:
in Exp2 was sterilized to eliminate microbial inuence on water or
g biomass (Y1 Y0)
P uptake by the plants. The sterilization process of Exp2 released APUE( )= (5)
some bioavailable nutrient into the soil, and therefore we reduced mg P fertilizer Pf
the N and P fertilizer dose. Furthermore, there were three harvest where: Y1 and Y0 are total biomass of P1 and P0 plants respectively;
times in Exp1 to follow plant water and P utilization dynamics but Pf is the rate of fertilizer P applied in P1 treatments.
only one harvest time in Exp2, which was at the same potato devel- Water soluble P (WSP) in the soil was measured at each har-
opment stage as the second harvest of Exp1. The fertilizer used in vest time in pooled supernatants after two sequential extractions
Exp1 was compound fertilizer while the fertilizer used in Exp2 was of 1 g of fresh soil shaken with 50 mL of deionized water for 1 h at
pure chemical. The soil used in both experiments had relatively low 20 C followed by centrifuging for 30 min at 10,733 g at 20 C and
concentration of P to induce P deciency in the plants. was analyzed with a spectrophotometer at 890 nm after the addi-
tion of vanadate molybdate (Sissingh, 1971). Microbial biomass P
2.2. Sampling, measurements, calculations and statistical (MBP) was measured at each harvest time by measuring the resin P
analyses difference with or without hexmo addition to soil. The resin P was
extracted with anion exchange membranes (AEM) where 2 g fresh
The irrigation volumes (Wi ) were calculated at each irrigation soil was suspended in 30 ml deionized water (Kouno et al., 1995).
time based on TDR-measurements with two to four replicates of Soil mineral N was only measured at nal harvest. The soil nitrate-N
each treatment using the formula: (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
W i (L) = Vs(L) ( p  a ) (1) (Bran + Luebbe, Germany). We assumed that P transport to root
where: Vs is the soil volume in each pot,  p is the volumetric water surface was the sum of mass ow and diffusion. Percentage of P
content (%) at pot water holding capacity and  a is the actual volu- transport to the root surface by mass ow (P transport massow ) and
metric water content (%) as measured by TDR. diffusion (P transport diffusion ) was calculated according to Scheffer
The applied water volume during the irrigation treatment was and Schachtschabel (1979) as:
the sum of the irrigation volumes (Wi ) at each irrigation time. Aver- Pm
Ptransportmassow = 100 (6)
age soil volumetric water content () was calculated by averaging Phyp
the soil water content before and after irrigation.
In Exp1, the rst, second and nal harvests were done after 0, 27
Ptransportdiffusion = 100 Ptransportmassow (7)
and 52 (nal harvest) days irrigation treatments. Leaf water poten-
tial was measured one day before nal harvest with a pressure where: Pm is the measured P concentrations which was taken as the
chamber (Soil Moisture Equipment Corp., Santa Barbara, CA, USA) bulk soil WSP before start of irrigation treatments converted to mg
on the second (from the top) fully expanded upper leaf. Shoots (leaf P/L soil solution according to Koopmans et al., (2010) and Blaauw
and stem) and underground parts (root and tuber) of the plant, bulk et al., (1988). Phyp is the calculated hypothetical P concentrations
and rhizosphere soil were collected at each harvest. At nal harvest, in the soil solution if plant P uptake was achieved solely by mass
leaf area was measured with a leaf area meter (LI-3100C, Li-Cor Lin- ow transport.
coln, NE, USA). The bulk soil was collected by mixing all the pot soil
(topsoil and subsoil) evenly. The rhizosphere soil was collected by Pm = 0.00375 Pw 1.466 (8)
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 Phyp (mg/L)
was recorded. Total (dry) biomass is the sum weight of roots, tubers,
plant P uptake
stems and leafs. Root to shoot ratio (RSR) was calculated as: = (9)
water used by transpiration during irrigation treatment
100 root dry biomass
RSR(%) = (2) where: Pw is the soil P concentration expressed by mg P2 O5 / L air
shoot dry biomass
dried soil.
Water use efciency (WUE g L1 ) was calculated as: The soil mineral N balance (SNmin ) was calculated as the differ-
ence between the sum of soil mineral N (SNt1 ) and applied mineral
WUE(g L1 ) N (AN) at the onset of the experiment (t1 ) and the sum of bulk soil
plant total biomass increment (g) mineral N (SNt2 ) and average plant biomass N (PN) at the time of
= (3) nal harvest (t2 ):
applied water volume during irrigation treatment (L)
SNmin = (SNt1 + AN)(SNt2 + PN)(10)
Plant samples were divided into shoots and underground parts,
and the dried samples were ground to a ne powder. Total P con- In Exp2, the harvest was done 30 days after irrigation treatments
centration was measured at each harvest time after ashing (450 C), started. Plant and bulk soil samples were collected. Plant biomass,
solubilizing in 13.9% hydrochloric acid and 21.7% nitric acid and WSP, total P concentration of plant, root to shoot ratio and P uptake
was analyzed using a spectrophotometer after the addition of were measured and calculated in the same way as exp1. Soil acid
vanadate molybdate (Stufns, 1967). Total nitrogen (N) concen- phosphomonoesterase (APM) activity was measured with a spec-
tration was measured at nal harvest with LECO CNS-1000 (LECO trophotometer at 420 nm (Tabatabai and Bremner, 1969) after 1 g
Corp., St. Joseph, MI. USA) according to ISO 13,878. Plant P and N (dry weight) fresh soil had been incubated in a universal buffer at
uptake (mg plant1 ) was calculated as total P and N concentration 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 Exp2 (Table 1), possibly due to the higher P-availability in the P0
hour. treatment of Exp2 caused by the soil sterilization. In both exper-
The statistical analyses were conducted using the MIXED pro- iments under P1, the total plant biomass under decit irrigation
cedure of SAS (9.3). The xed effects were irrigation treatments, P (DI) and partial root zone drying irrigation (PRD) treatments was
fertilization levels and irrigation treatments P fertilization levels. similar, and was about 24% lower than that of full irrigation (FI).
Replicate was included as a random effect. The correlation analysis However, at P0 in Exp1, no difference of plant biomass was found
was done by the CORR procedure of SAS (9.3). for the three irrigation regimes, giving rise to an interactive effect
between P and irrigation at nal harvest, while in Exp2, the total
3. Results 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
3.1. Plant growth, soil water dynamics and water use efciency both experiments (Fig. 2). Further, the root biomass was affected by
under different irrigation regimes irrigation treatments but differently between the two experiments
as root biomass in Exp1 was lower under DI and PRD compared to
The application of phosphorus (P) fertilizer increased potato FI, while root biomass in Exp2 was higher under DI and FI compared
growth in both experiments, but to a larger extent in Exp1 than 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 P had similar dynamic as WSP, but the value was 4-5 times larger;
decreased during plant growth. Under P1, P concentration of shoots data not shown). MBP also had the tendency to increase along with
initially was higher under FI than DI and PRD, later became almost the growth of potato (Fig. 6; Exp1) and MBP in rhizosphere of DI
equal to DI and PRD, and nally decreased and became lower than was larger than FI and PRD. APM concentration of PRD was higher
DI and PRD when harvested at 52 day. At P0 levels in Exp1, shoot than DI and also tended to be higher than FI at both P levels (Fig. 6;
P concentration of PRD was higher than FI and DI when harvested Exp2).
at 52 day, while PRD had similar biomass production as DI. In both Agronomic P use efciency (APUE g biomass/g P applied per
experiments at P1 levels, DI and PRD decreased P uptake by about plant) was signicantly inuenced by irrigation treatments at nal
22% compared with FI at nal harvest, and P uptake of DI and PRD harvest in Exp1. The APUE (g biomass/g P applied per plant) of FI, DI
was similar. All the three irrigation treatments had similar P uptake and PRD were 107, 60 and 67, respectively. The APUE of FI treatment
at P0 levels when harvested at 52 day in Exp1, giving rise to a signif- was much larger than DI and PRD, while APUE of PRD was slightly
icant interaction between P and irrigation treatments. In contrast, P higher than DI. Generally, soil P transport to the root surface was
uptake in Exp2 at P0 and P1 levels showed the same pattern for the dominated by diffusion, which accounted for more than 95% of the
three irrigation treatments with DI and PRD being lower than FI. total P uptake (Fig. 7). The application of P fertilizer decreased the
P concentration and plant uptake were higher at P0 levels in Exp2 proportion of P transported to the root surface by diffusion. The
than in Exp1 as mentioned earlier possibly due to the soil steriliza- proportion of P transported to the root surface by diffusion was
tion in Exp2. The P uptake positively correlated to plant root dry similar under DI and PRD and higher than under FI (Fig 7).
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, 3.3. Effect of irrigation on plant N uptake, plant N:P ratio and soil
P < 0.001) (Fig. 5). mineral N balance
The application of P fertilizer increased water soluble P (WSP)
and microbial biomass P (MBP) but decreased activity of acid phos- The application of P fertilizer strongly increased potato N uptake
phomonoesterase (APM) (Fig. 6). WSP in rhizosphere soil decreased but decreased plant N:P ratio (Table 3). Shoot and total N uptake
along with potato growth at P1 levels, while it was much lower and under PRD was higher than under DI at P1 and also tended to be
almost constant at the P0 level. WSP in bulk soil showed similar higher than DI at P0. While shoot and total N uptake were highest
dynamics as in rhizosphere soil but with smaller values (data not under FI at P1, this was strikingly different at P0 levels where PRD
shown). When harvested at 27 (Exp1) and 30 day (Exp2), WSP of had the highest N uptake, and therefore there was a signicant
DI was higher than PRD in both rhizosphere and bulk soil. When interaction between P levels and irrigation treatment. Shoot and
harvested at 52 day, WSP of FI was lower than DI and PRD (resin 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 The NH4 -N concentrations of rhizosphere and bulk soil resembled
with declining soil water content at both high and low P levels those of NO3 -N but DI was higher than PRD (P = 0.059). At the begin-
(curve not shown). ning of the experiment, NO3 -N and NH4 -N were 145 mg kg1 and
It was clear that P fertilizer decreased mineral N left in soil 132 mg kg1 soil, respectively. After harvest, NO3 -N had decreased
(Table 4). Soil NO3 -N concentration in PRD and DI was similar and to 235 mg kg1 and NH4 -N had decreased to 03 mg kg1 .
was higher than in FI in both rhizosphere and bulk soil. For the The application of P fertilizer also strongly decreased the soil
P0 levels, higher N uptake of PRD compared to FI was observed. 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


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
Fig. 7. Percentage of P transport to root surface by diffusion in total P transport (%) as
P-uptake were small and non-signicant between irrigation treat-
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 ments.
PRD: partial root zone drying irrigation). Error bars indicate SE (n = 6). Signicant Plant P uptake and P use efciency may be improved by: (i) roots
differences are indicated by letters: grouped columns without letters in common exploring a larger soil volume, (ii) easing transport of P from bulk
are signicantly different between irrigation treatments at P < 0.05. Symbol [] soil to root surface and (iii) transforming non-available P forms into
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
plant-available P (Rengel, 2008). In the current study, the P fertilizer
were signicant interactions of P fertilization levels and irrigation methods at 27 was applied only to the top 2022 cm soil in both experiments. As
and 52 days harvests. the P easily binds to soil particles, downward movement of P under
these experimental conditions would be small, and the fertilizer
FI at P0 levels where it amounted to 46% of the N fertilizer added at P would be concentrated in the top soil layer; the same situation
the start of the experiment, but at P1 levels it was lowest under FI as usually found under eld conditions. The factors that affect P
and thus an interactive effect of P and irrigation was found. At both uptake i.e. root development and function, P diffusion in soil and
P levels however, SNmin was larger under DI than under PRD. plant available P, are discussed in the following paragraphs.
Root architecture plays an important role in P acquisition
4. Discussion (Gahoonia, 1994; Lynch, 1995) because it determines the size of
the explored volume of soil. The repressed primary root growth
It was hypothesized that partial root zone drying irrigation in response to P deciency conditions is based on the presence of
(PRD) can improve nitrogen (N) uptake, and the improved N uptake NO3 (Medici et al., 2015), clearly indicating an integrated response
might be accompanied by an improved phosphorus (P) uptake. of root architecture formation to nitrate and phosphate. Consistent
The result showed that compared to DI, PRD improved N uptake. with Hodge, (2004), an enhanced root growth due to P fertiliza-
However, we did not nd improved P uptake under PRD, instead 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. activity. A 10% reduction of soil moisture may decrease acid phos-
Although no root mass difference was found in previous studies in phatase activity by 3040% (Sardans and Penuelas, 2005). Acid
potato irrigated with PRD, DI and FI (Ahmadi et al., 2011). FI had the phosphomonoesterase activity was only measured in Exp2 with
highest root mass and PRD restricted root growth compared with sterile soil in this study, so it was assumed that they were secreted
DI (only in Exp2). An increased root to shoot ratio was found under by plant roots. The results showed that acid phosphatase activity
P0, which is a typical response to P decient condition (Wissuwa was decreased by DI treatments compared to FI (Fig. 6), but PRD
et al., 2005), as preferential assimilate distribution to the roots treatment had root acid phosphatase activity similar to FI, which
potentially may increase the plant P uptake. The pots used in the were higher than DI. This nding differed from results of previ-
experiment were 16 cm in diameter, which could have affected the ous work (Li et al., 2010) where PRD decreased the activity of acid
measured responses with respect to plant biomass, root biomass phosphatase.
etc. Compared with plants grown in the eld, the potted plants had In a review of 40 fertilization studies it was concluded that, N:P
less soil volume for root growth. Also, the arrangement of plants in ratios in plants >16 are indicative of P limitation, while N: P < 14
the greenhouse left more space for top growth, which may affect indicate N limitation. Ratios of N:P between 14 and 16 indicate
plant shape, light interception and transpiration rate. As a result, either N or P or possibly N and P together are limiting plant growth
the total biomass, root biomass, N uptake, P uptake and tuber pro- (Koerselman and Meuleman, 1996). The N:P ratios reect the rel-
duction might be different from plants grown in eld. The effects ative availability of N and P to plants and may indicate the degree
on root distribution and root length density might have affected P of N or P deciency experienced by a plant (Gusewell et al., 2003).
uptake (Miguel et al., 2015), and therefore further experiments are According to this standard, as can been seen from Table 3, all the
needed to verify their role on P uptake. Diffusion, the main pro- irrigation treatments at the low P levels suffered a severe P de-
cess making P available to roots, supplies P from the bulk soil to ciency, while treatments at the high P levels under FI suffered slight
the root surface (Comerford, 1998). Current results showed that, N deciency, and PRD suffered slight P deciency. Shoot N:P ratio
diffusion transferred more than 95% of P from soil to roots. There- also indicated that PRD treatments went through a more severe
fore, no matter the kind of irrigation treatment used, diffusion was P deciency than DI and FI and this deciency might have trig-
the main way of P transport to the root surface, which implied gered more root phosphatase secretion in PRD than in DI. The higher
that the percentage of P transport to the root surface by diffusion phosphatase activity of PRD might have resulted in a higher min-
was less affected by irrigation pattern than by soil water content. eralization of Po under PRD than DI. However this Po contribution
Plant growth is also an important factor affecting P uptake (White to plant total P uptake was not enough to result in a difference
et al., 2005b) as enhanced root growth will increase the uptake of P uptake under DI and PRD, since WSP tended to be higher in
by increasing the explored soil volume and cross sectional area of DI and ultimately similar P uptake was achieved under DI and
diffusion. So any other environmental factors that improve root PRD. The trend that shoot N:P ratio increased with declining soil
growth may also be accompanied with a higher P uptake. water content was in agreement with Cernusak et al. (2010) and
Irrigation using the PRD method is accompanied by drying and was explained by differential changes in the mobility of nitrate vs
rewetting cycles (D/RW) of the soil prole. D/RW causes microbial phosphate in response to declining soil water content.
death and cell lysis (Turner et al., 2003), followed by an increase In the present study, we found improved N uptake of PRD both
in soil respiration rate indicating enhanced microbial activity and at low and high P levels compared to DI. The soil mineral N balance
ushes of mineral N and P into soil solution (Birch 1958; Xiang et al., (SNmin ) as dened here measures the decrease in soil mineral N
2008; Butterly et al., 2009). However, other results have indicated from planting to harvest other than explained by plant N uptake.
that soil available phosphate is not affected by wetting a dry soil As SNmin was positive, there was a reduction of mineral N rather
(Cui and Caldwell, 1997). The soil microbial biomass plays a very than soil N mineralization as found in the study by Sun et al. (2013).
important role in soil fertility both as a source and sink for mineral The possible reasons why SNmin was positive in PRD, DI and FI
nutrients (Kouno et al., 1995). In the present study, D/RW of PRD were either denitrication, ammonia volatilization or immobiliza-
caused a reduction in the size of microbial biomass compared to DI, tion through microbial conversion of N to organic forms (Cameron
measured as biomass P which is consistent with previous studies et al., 2013). Of these, immobilization of N to forms which are tem-
(Wu and Brookes, 2005; Gordon et al., 2008). The choice of dry- porarily unavailable to plants was the most probable cause for the
ing to 8% (Vol) was based on the soil water characteristics curve different SNmin under DI and PRD, since denitrication was likely
and results from the article of Liu et al., (2008) suggesting that the low due to the soil water content being kept at or below 70% eld
dry side should be allowed to dry to a water potential of around capacity as well as the low soil pH (below 7) was not favoring
80 kPa to induce relevant ABA production in potatoes. In current NH3 volatilization. There are substantial indirect and direct evi-
experiments, except from reduced size of microbial biomass, there dence supporting that the competition exists between plants and
was additional evidence of severe stress caused by D/RW of PRD as microorganisms (Kaye and Hart, 1997) and the applied inorganic N
indicated by lower leaf area, total biomass, root biomass and leaf can be rapidly taken up by the soil microbes (Nannipieri et al., 1990).
water potential, all of which implied that allowing the water con- Up to 50% immobilization of fertilizer N has also been found in other
tent to decrease to 8% (Vol) for the dry side of PRD before switching pot and eld studies (Blankenau et al., 2000) and in the present
between the sides was too extreme to maintain growth. However, experiments seemed to be favored by the high concentration of
no ush of plant available P under PRD was found in this study com- mineral N in the soil under P0 conditions where plant uptake was
pared with DI. When given same amount of P fertilizer, WSP, which limited by P deciency. Also the use of ammonium nitrate as fer-
is plant available, decreased along with plant P uptake. As a result, tilizer may have favored immobilization processes as ammonium
it was inappropriate to compare plant available P in case the plant seems to be the preferred substrate for bacterial immobilization
P uptake was different. Higher P availability (soil WSP and resin (Wickramasinghe et al., 1985; Blankenau et al., 2002). The lower
P) was found under DI instead of PRD with high P levels at 27 day SNmin found under PRD than under DI in P1 may also suggest less
(Exp1) and 30 day (Exp2) harvest, which may indicate a xation of N immobilization in PRD and this corresponded to the higher plant
available P by D/RW (Fig. 6). N uptake under PRD. The tendency of larger root biomass especially
Organic P (Po) accounts for 5070% of the total P in soils (Borie under P1 which can provide carbon for microbial metabolism also
and Rubio, 2003). Phosphatase catalyzing the hydrolysis of Po is corresponded to the tendency of higher MBP in DI than PRD pots.
important to soil P availability. Soil moisture affects the soil microbe The estimation of mineral N immobilization under DI and PRD at
and root activity and is an important factor affecting soil enzyme P1 tted the value of SNmin by the calculation of soil microbial
C. Liu et al. / Agricultural Water Management 159 (2015) 6676 75

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