Agricultural Water Management 123 (2013) 65–70

Contents lists available at SciVerse ScienceDirect

Agricultural Water Management

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

Effect of partial root-zone drying irrigation timing on potato tuber

yield and water use efficiency
Wendy Yactayo, David A. Ramírez ∗ , Raymundo Gutiérrez, Víctor Mares,
Adolfo Posadas, Roberto Quiroz
International Potato Center (CIP), Apartado 1558, Lima 12, Peru

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

Article history: Partial root-zone drying (PRD) is an irrigation technique which has shown increased water use efficiency
Received 2 January 2013 (WUE) without yield reductions in potato and other crops. However, questions remain as to the effect
Accepted 13 March 2013 of the water restriction initiation timing and the level of alternate water restriction on the response of
Available online 16 April 2013
the potato crop. In this study, we tested: two PRD treatments with 25% (PRD25 ) and 50% (PRD50 ) of total
water used in full irrigation (FI, as control), and a deficit irrigation treatment with 50% of water restriction
Keywords:
(DI50 ). Two water restriction initiation timings were tested at: 6 weeks (WRIT6w) and 8 weeks (WRIT8w)
Osmotic adjustment
after planting. Osmotic potential (␲), osmotic adjustment, relative water content and chlorophyll con-
PRD
Stay green effect
centration were assessed in four dates during the growing period. PRD50 initiated at WRIT6w showed the
Solanum tuberosum highest WUE without a tuber yield reduction respect to the control. While plants under PRDs and DI50
showed lower ␲ than FI, PRDs treatments promoted higher osmotic adjustment particularly in WRIT6w.
Our study suggests that early PRDs with mild water restriction allow drought hardiness (improving water
stress response) and water saving avoiding a dramatic yield tuber reduction.
© 2013 Elsevier B.V. All rights reserved.

1. Introduction irrigated water below the maximum crop evapotranspiration) and

The potato (Solanum tuberosum L.) cropping area has been
expanding in developing countries, particularly in Asia and Africa
(Haverkort et al., 2004). This expansion is opening new cropping
areas in some environments that could be negatively affected by
global warming (Thiele et al., 2010). A reduction of 18–32% of
potato global yield caused by climate change is projected to occur
during 2010–2039 (Hijmans, 2003). Alongside temperature and
atmospheric CO2 increases, climate change will bring about a larger
rainfall variability and evaporative demand (Kundzewicz et al.,
2007), a situation that will affect potato, which is a drought sen-
sible crop (van Loon, 1981). Furthermore, competing water use
for agriculture and other activities is becoming an important issue
worldwide (Naylor, 2009). Under these unpredictable and highly
variable conditions, increasing water use efficiency (WUE, tuber
yield per amount of water applied) by irrigated potato crops is an
important objective.
Irrigation water saving techniques have not been investigated
in potato to the same extent as in other crops, particularly cere-
als and fruit trees (Kang and Zhang, 2004). However, work so far
conducted in irrigated potato indicates that deficit irrigation (DI,
∗ Corresponding author. Tel.: +51 1 3175312; fax: +51 1 317 5329.
E-mail address: d.ramirez@cgiar.org (D.A. Ramírez).
partial root-zone drying (PRD, alternated irrigation of the root-zone
by watering of one furrow and keeping dry the adjacent one until
the next watering cycle) are two promising irrigation techniques to
save water with a concomitant WUE increase and no yield reduc-
tion (Liu et al., 2006a,b; Jensen et al., 2010; Xie et al., 2012). In
several experiments, PRD has given better results than DI and full
irrigation, allowing for 39–50% of water saving, while increasing
WUE without significant tuber yield reduction (Liu et al., 2006a;
Saeed et al., 2008; Shahnazari et al., 2008; Jovanovic et al., 2010; Xie
et al., 2012). Higher marketable tuber size, soil N-availability and
antioxidants in tubers have been obtained in PRD trials in potato
(Rojas et al., 2007; Shahnazari et al., 2007; Shahnazari et al., 2008;
Jovanovic et al., 2010). Notwithstanding these promising results
and in order to maximize the benefits of the technique, issues such
as the best combination of initiation timing, duration, and intensity
of water restriction (sensu Jefferies, 1995) remain to be addressed.
It is also important to assess how water restriction trials could con-
tribute to enhance the expression of drought tolerant traits without
tuber yield penalization. Saeed et al. (2008) and Xu et al. (2011)
obtained higher tuber yield in PRD treatments when water restric-
tion was initiated soon after tuber initiation, both in pot and in
field trials, respectively. Although these experiments tested differ-
ent initiation timing and duration of water restriction, the amount
of water applied was similar across treatments. On the other hand,
PRD trials with different levels of water restriction (e.g. Xie et al.,

0378-3774/$ – see front matter © 2013 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.agwat.2013.03.009
66 W. Yactayo et al. / Agricultural Water Management 123 (2013) 65–70
2012) have not considered the assessment of the initiation timing
and duration of the restriction.
Osmotic adjustment (accumulation of low-molecular organic
substances into the cell that promote the osmotic influx of water;
Larcher, 2003) and “stay green” (delayed senescence) are some of
the desirable traits in the selection programs for drought resis-
tance in crops (Blum, 2011). The expression of both traits have been
enhanced using different PRD treatments in potato (Xu et al., 2011;
Jensen et al., 2010) which indicates that an appropriate combina-
tion of initiation timing, duration and intensity of water restriction
would induce the activation of these traits with a concomitant WUE
rise (see Xu et al., 2011). In this paper we report the results of PRD
trials with different initiation timing, duration and levels of water
restriction under field conditions in an arid region with no rain,
and thus without rainfall confounding effect. The aims of the work
were: i) to assess WUE under contrasted PRD treatments with dif-
ferent water restriction initiation timing, ii) to determine whether
the PRD treatment induced water stress tolerance as indicated by
osmotic adjustment and “stay green” effects.
2. Materials and Methods
2.1. Experimental site
The study was carried out at the International Potato Cen-
ter (CIP) experimental station in Lima, Peru (12.08◦ W; 76.95◦ S,
244 m asl) from June to November 2010. The station is located
in the Peruvian desert with average yearly precipitation, maxi-
mum − minimum temperatures, atmospheric humidity and global
radiation of 23 mm, 22.4–16.5 ◦ C, 79.7% and 14.1 MJ m−2 d−1 ,
respectively (2008–2010, CIP Meteorological Station). The soil is
sandy loam (50, 32 and 18% of sand, lime and clay, respectively),
with an organic matter content, field capacity, bulk density, pH and
electric conductivity of 8%, 14.6%, 1.4 g cm−3 , 7.6 and 2.4 dS m−1 ,
respectively (Laboratorio de Análisis de Suelo, Plantas, Aguas y
Fertilizantes–Universidad Nacional Agraria La Molina, Lima, Peru).
2.2. Description and design of experiment
The potato variety tested was UNICA (CIP code N◦ 392797.22)
which is an early variety and considered to be tolerant to viruses
and high temperature (Gutiérrez et al., 2007). The fertilizer appli-
cation consisted of 200:140:160 kg ha−1 as N:P2 O5 :K2 O. One half
of the N was supplied at planting and the rest at hilling. The phy-
tosanitary control against pests and diseases was carried out every
20 days.
The experimental unit consisted of a 4.5 × 4.5 m2 plot divided
into 6 rows (0.9 m apart from each other) and 5 ridges. Each plot
contained 65 plants at a distance of 0.3 m. The evaluations were
conducted on 13 plants from the center of the plot to avoid border
effects. The irrigation treatments were: full irrigation (FI), which
supplied 100% of the crop water demand, deficit irrigation (DI),
amounting to 50% of FI, partial root-zone drying (PRD) which sup-
plied 50 and 25% of FI (PRD50 and PRD25 , respectively). The furrow
irrigation schedule for all treatments was based on infiltration
assessments using the method of cylinder infiltrometer (FAO, 2012)
fitting the following power function:
WI = 0.24 T 0.77
where WI is infiltrated water layer (mm) and T is irrigation time
(min). Each 12 days the gravimetric soil water content () of the
plots was assessed at 0.25 m depths and the crop water demand
(from evapotranspiration and percolation) was calculated by sub-
tracting field capacity minus . The furrow irrigation time, required
to compensate crop water demand, was calculated from (1). Water-
ing was performed using siphons to obtain similar water flow into
the furrows. In order to test the effect of different water restric-
tion initiation timing (WRIT) the irrigation treatments began at 6
(WRIT6w) and 8 (WRIT8w) weeks after planting. The irrigation fre-
quency was every 12 days (with a respective shift of furrows in
the PRD treatments) with the following total number of irrigation
events from planting to harvest: 3 and 4 before the initiation of the
different irrigation treatments for WRIT6w and WRIT8w, respec-
tively; 6 and 5 during the experimental phase for WRIT6w and
WRIT8w, respectively. The length of the cropping period was 17
weeks with 10 and 8 weeks of experimental phase for WRIT6w and
WRIT8w, respectively.
2.3. Ecophysiological and agronomic measurements
All the ecophysiological measurements were carried out in four
occasions during the experimental period in samples taken from
three central rows within the plot. Two plants per plot were sam-
pled and one leaflet from an expanded and sun-exposed leaf located
in the upper third section of the plant canopy collected between
6:00 and 7:00 am local time and their fresh weight (FW) were
recorded. The samples were immersed in distilled water during 6 h
at 6 ◦ C and weighted again (saturated weight, SW). Leaflets were
then dried during 48 h at 80 ◦ C and weighted (dry weight, DW).
Relative water content (RWC) was estimated as:
RWC = (FW − DW ) ∗ 100/(SW − DW ) (2)
Osmotic potential () was measured using Turner’s (1981) pro-
tocol which involves the rupture of the cell membrane brought
about by an abrupt leaf defrosting which causes that water poten-
tial equals . A leaflet circular sample (0.5 cm of diameter) was
immersed in liquid nitrogen and after that conserved at −80 ◦ C. The
water potential was measured in defrosted samples using a dew
point potentiometer (Wescor, Logan, UT, USA). Linear regressions
were fitted through the value pairs (RWC, ). Osmotic adjustment
was estimated as the slope (RWC/␲) of the fitted functions
where low slope values were interpreted as high osmotic adjust-
ment and vice versa (Morgan, 1983, 1992).
Chlorophyll content was estimated with a portable chlorophyll
meter (SPAD-502 model, Konica Minolta, Sakai, Osaka, Japan). Nine
readings per leaf were averaged per plant.
Tuber yield (kg ha−1 ) was estimated from tuber fresh weight
produced by the plants harvested from the three central rows of the
plots. Irrigation water use efficiency (WUE, kg m−3 ) was calculated
as the ratio between tuber yield and total WI.
2.4. Statistical analysis
Main irrigation treatment effects and interactions were assessed
by ANOVA using a randomized block design. For comparing water
restriction treatments against the control (FI) the Dunnett MRT
test was applied. Furthermore, Duncan’s test was also applied to
the data in order to assess differences among irrigation treatments
in both early and late irrigation onset timing. Ecophysiological
measurements taken sequentially over time introduce carry over
effects, i.e. observations close in time may be more related than
observations far apart in time. Repeated measurements analyses, as
described by Wolfinger and Chang (1998), was used as a corrective
measure. The analysis of variance generated by the General Lin-
(1)
eal Model (GLM) allows the determination of statistical differences
between treatments, as a function of time, with a pre-established
probability level (P value) of 5%. A significant difference indicates
that, on the one hand, the variation due to treatment, at a particular
time within the experiment, was greater than the variation among
plants within each treatment. On the other hand, it indicates that
the number of replicates was enough to reach a robust assessment
of the differences between plants submitted to different irrigation
 W. Yactayo et al. / Agricultural Water Management 123 (2013) 65–70 67

Table 1
F-value of ANOVA of tuber yield and irrigation water use efficiency (WUE). Water
restriction initiation timing at 6 (WRIT6w) and 8 (WRIT8w) weeks after planting.
IT = irrigation treatment.

WRIT6w WRIT8w

IT Block IT Block
** ** *
Tuber yield 19.7 11.9 6.2 22.5**
WUE 11.8** 13. 0** 4.3* 18.2**
**
P < 0.01,
*
P < 0.05.

treatments. The significance of the difference between the slopes of

the linear regression between RWC vs  was tested by the t-Student
following the verification of the homogeneity of the variance. All
the statistical analyses were run with SAS v8.02 software (SAS Insti-
tute, Cary, NC, USA).

3. Results

3.1. Total crop water demand, tuber yield and water use efficiency

Irrigation treatments and blocks effects were significant in WUE

and tuber yield (Table 1). Total crop water demand was higher
when the restricted irrigation was initiated 8 weeks after planting
(WRIT8w; Table 2) as compared to WRIT6w. The water quantity
saved, when restricted irrigation was initiated at WRIT6w respect
to WRIT8w, was 99.1, 68.2, and 55.6 m3 ha−1 in PRD25 , DI50 and Fig. 1. Average ± standard error (n = 4) of tuber yield (a) and water use efficiency
PRD50 , respectively. The reduction (%) order of crop water demand (WUE) (b) in irrigation treatments (FI = full irrigation, DI50 = deficit irrigation with
with respect to FI was PRD25 > DI50 > PRD50 in both WRITs (Table 2). 50% of FI total water, PRD50 and PRD25 = partial root-zone drying with 50 and 25%
of FI total water, respectively) at two irrigation onset timing: 6 (WRIT6w) and
The restricted irrigation treatments DI50 and PRD25 significantly 8 (WRIT8w) weeks after planting. Different letters indicate differences (P < 0.05)
reduced (P < 0.05) tuber yield compared with FI, regardless of WRIT detected by Duncan’s multi-treatments comparison test.
(Fig. 1a). In contrast, no differences in tuber yield relative to FI were
observed by the PRD50 treatment (Fig. 1a). As to WUE, it was sig-
−2.1 to −2.7 MPa for WRIT 2 (Fig. 2d). For WRIT6w, significant
nificantly higher under PRD50 , compared with all other treatments,
differences between irrigation treatments in the first and third
when water restriction was initiated at the earlier date, WRIT6w,
assessment were found (Fig. 2c), being PRD25 the treatment show-
but no differences among watering treatments were observed with
ing the lowest  values. For WRIT8w, significant differences in
WRIT8w (Fig. 1b). In the WRIT6w situation, the significant increase
 between treatment were evident at the first, third and fourth
in WUE brought about by PRD50 amounted to 58% with respect to
assessment (Fig. 2d). ChlSPAD values fell from 48.8 to 37.4 SPAD
FI.
units during WRIT6w (Fig. 2e) and from 49.0 to 39.0 SPAD units
during WRIT8w (Fig. 2f). FI showed significant lowest values of
3.2. Physiological responses
ChlSPAD in all the assessments in WRIT6w and at the last assess-
ment in WRIT8w (Fig. 2e, f). Actually, Irrigation Treatment x Time
All assessed variables showed significant differences between
effect was the only significant interaction for ChlSPAD.
repeated measurements along time after the date water restric-
At WRIT6w, equations and determination coefficients
tion treatments were initiated. Although no significant effect of
obtained were, FI: Y = 13.4X + 112.9, r2 = 0.84; DI50 :
irrigation treatments on RWC were found (Table 3), RWC signifi-
Y = 11.6X + 109.4, r = 0.92; PRD50 : Y = 10.0X + 108.0, r2 = 0.95;
2
cantly decreased along the experimental phase, both at WRIT6w
PRD25 : Y = 10.9X + 108.6, r2 = 0.96 (all F-values of the linear regres-
(Fig. 2a) and WRIT8w (Fig. 2b). Significant irrigation treatments
sion analysis were significant at P < 0.05). FI and PRD50 showed
effects on both osmotic potential (␲) and chlorophyll content
the highest (13.4 ± 4.2) and lowest slope (10.0 ± 1.5), respectively,
(ChlSPAD ) were observed (Table 3), regardless of WRIT. With respect
to time effects, ␲ showed a consistent drop from WRIT onwards.
It dropped from −2.1 to −2.8 MPa for WRIT6w (Fig. 2c) and from Table 3
F-values of ANOVA with repeated measurements in time corresponding to leaflet
relative water content (RWC), osmotic potential () and chlorophyll concentration
Table 2
(ChlSPAD ) obtained at two water restriction initiation timing: 6 (WRIT6w) and 8
Average of total crop water demand (CWD, m3 ha−1 ) in irrigation treatments at two
(WRIT8w) weeks after planting.
water restriction initiation timing: 6 (WRIT6w) and 8 (WRIT8w) weeks after plant-
ing. The percentage reduction of CWD was estimated with respect to full irrigation Irrigation Time Block IT × time
treatment (FI, control). DI50 = deficit irrigation treatment with 50% of FI total water, treatment (IT)
PRD25 and PRD50 = partial root-zone drying treatments with 25 and 50% of FI total
water respectively. WRIT6w RWC 0.29 n.s. 29.1** 0.58 n.s. 0.96 n.s.
 5.92* 75.9** 1.30 n.s. 1.83 n.s.
WRIT6w WRIT8w ChlSPAD 51.7** 500.2** 7.00* 21.6**
WRIT8w RWC 0.16n.s. 19.3** 0.62 n.s. 0.66 n.s.
CWD Reduction (%) CWD Reduction (%)  3.93* 56.5** 0.31 n.s. 1.48 n.s.
FI 2225.1 2218.7 ChlSPAD 21.78** 177.0** 3.90* 18.0**
DI50 1375.7 38 1443.9 35 n.s.: no significant (P > 0.05)
PRD50 1519.8 32 1575.4 29 **
P < 0.01.
PRD25 1023.1 54 1122.2 49 *
P < 0.05.
68 W. Yactayo et al. / Agricultural Water Management 123 (2013) 65–70

Fig. 2. Average ± standard error of chlorophyll concentration (ChlSPAD ) (a,b), leaflet relative water content (RWC) (c,d) and osmotic potential () (e,f) under irrigation
treatments (FI = full irrigation, DI50 = deficit irrigation with 50% of FI total water, PRD50 and PRD25 = partial root-zone drying with 50 and 25% of FI total water, respectively) at
two water restriction initiation timing: 6 (WRIT6w) and 8 (WRIT8w) weeks after planting. ** P < 0.01, *P < 0.05, n.s. = no significant (P > 0.05). DAT = days after treatment.

in the linear regression analysis between RWC vs  (Fig. 3a);
however, there was no significant differences among slopes during
WRIT6w (Table 4). At WRIT8w, equations and determination
coefficients obtained were, FI: Y = 22.2X + 134.7, r2 = 0.84; DI50 :
Y = 21.8X + 132.0, r2 = 0.97; PRD50 : Y = 15.1X + 121.0, r2 = 0.98;
PRD25 : Y = 12.3X + 107.2, r2 = 0.92 (all F-values of the linear regres-
sion analysis were significant at P < 0.05). The highest and lowest
slope of the regression between RWC vs  corresponded to FI
(22.2 ± 5.41) and PRD25 (12.3 ± 2.6) respectively (Fig. 3b). Only
marginal significant differences (P = 0.07) were found between DI50
and PRD25 slopes during WRIT8w. We found that slopes in WRIT6w
were marginally significant lower (at P < 0.1) than WRIT8w slopes
(higher osmotic adjustment) only in PRD50 (P = 0.09) and DI50
(P = 0.06) treatments.
4. Discussion
4.1. WRIT and level of water restriction effects on tuber yield and
WUE
The earliest PRD50 treatment caused the highest WUE while
maintaining tuber yield relative to the control. Our results suggest
that the combination of the watering technique (PRD vs DI), level of
water restriction and initiation timing are critical factors determin-
ing the yield and WUE response of the potato crop. This statement is
supported by the fact that the PRD50 treatment showed higher WUE
and tuber yield than the DI50 and FI (without significant differences
in tuber yield) treatments, a result that coincides with published
reports (Liu et al., 2006a; Saeed et al., 2008; Shahnazari et al., 2008;
Jovanovic et al., 2010; Xie et al., 2012) but was only found in the
early irrigation onset timing.
The findings also suggest that the early PRD50 treatment allowed
for a tradeoff between water saving, with a concomitant WUE
increase, and the expression of some water stress tolerant mech-
anisms (see below) that maintained tuber yield. PRD has been
regarded as a mild, false and partial drought (Xu et al., 2011) which
agrees with Tardieu (2012) who highlights that “mild water deficits
are the common in agricultural conditions”. It is likely that this sim-
ulation of a drought event brought about by the PRD treatments
drives an increase in root development and density compared to
fixed deficit irrigation or conventional furrow irrigation in potato
(Kang and Zhang, 2004) which could enhance water and nutrient
uptake with a consequent maintenance of tuber yield. A highest
WUE in PRD trials in potato has been attributed to an improvement
 W. Yactayo et al. / Agricultural Water Management 123 (2013) 65–70
Fig. 3. Relationship between leaflet relative water content (RWC) and osmotic
potential (␲) under irrigation treatments (FI = full irrigation, DI50 = deficit irrigation
with 50% of FI total water, PRD50 and PRD25 = partial root-zone drying with 50 and
25% of FI total water, respectively) at two water restriction initiation timing (WRIT):
6 (a) and 8 (b) weeks after planting.
of light use efficiency driven by a leaf index reduction (Shahnazari
et al., 2007). However, we have not measured this variable to test
this hypothesis.
Our results are in agreement with Saeed et al. (2008) study
in potted potatoes, who recommended an WRIT of 6 weeks after
planting in a PRD trial which did not reduce tuber yield, allowing
for 21% of water saving and a WUE increment of 19%. Under field
conditions, Xu et al. (2011) found that an early PRD100 treatment
applied for a 6 weeks period followed by a subsequent return to
FI showed a highest tuber yield than the control and a late PRD
treatment.
4.2. Physiological responses to irrigation treatments
The observed absence of effects of irrigation treatments on RWC
coincides with Saeed et al. (2008) findings. As leaf growth requires
(among other factors) cell turgence (Larcher, 2003), it is likely that
the plant is able to maintain RWC at different levels of water restric-
tion whereas the RWC fall along time could reflect the reduction of
leaf elongation and growth with time.
Concerning , our results agree with Xu et al. (2011) who found
lower  (more negative) values in PRD treatments compared with
FI in a field trial in potato, which were attributed to a higher cell
solutes concentration. Since osmotic adjustment is an important
mechanism of water stress tolerance in crops (Blum, 2011), the
trend of a higher osmotic adjustment (lower slope of RWC vs 
fitted line) found in PRD treatments with respect to DI and FI,
suggests that PRD improves water stress response in potato. How-
ever because we found no significant differences likely caused by
our lower statistic power (n = 4), this hypothesis must be tested
in future trials. Moreover, an early PRD could promote a hardiness
effect (sensu Bruce et al., 2007) as suggested by our observation that
osmotic adjustment (slope) was higher in WRIT6w than WRIT8w
especially in PRD50 and DI50 treatments. Our finding is supported
69
by Xu et al.’s (2011) study, which suggests that early PRD enhances
xerophytic responses via hardening, improving drought tolerance
mechanisms without penalization in tuber yield.
“Stay green” or delayed senescence effect has been observed in
crops submitted to water stress and this mechanism was consid-
ered as the cause of productivity increases driven by the persistency
of photosynthesis brought about by water restriction (Thomas and
Howarth, 2000). Our results suggest that an early PRD applica-
tion (ca tuber initiation onset) in combination with medium water
restriction levels enhance WUE and maintain yield and that these
effects would be associated to improved osmotic adjustment and
“stay green” effects. In potato, xylem-ABA production has been sug-
gested as a major driver of enhanced WUE under induced water
restriction (Liu et al., 2006a). It has been suggested that ABA limits
ethylene production with a concomitant effect of senescence retar-
dation in leaves, flowers and fruits (Morgan and Drew, 1997; Sharp,
2002). In our work, the higher chlorophyll concentration caused by
PRD25 and DI suggests a large effect of water restriction on the
onset and process of leaf senescence, particularly by the delayed
initiation (WRIT8w) of differential watering. However, similar dif-
ferences were not observed as a consequence of WRIT6w, which
suggests that the relationship between water restriction and its
timing is not a simple one.
5. Conclusions
A challenge for the application of the PRD irrigation technique
to the potato crop is to establish the most appropriate combination
of the initiation timing, duration and intensity of water restriction,
and to understand the mechanisms that support water stress tol-
erance without yield tuber reduction. In this study we found that
an early PRD, initiated 6 weeks after planting, with a watering level
equivalent to 50% of full irrigation, increased WUE with no yield
reduction relative to full irrigation. Our results suggest that early
PRD improve water restriction tolerance in potato through a hard-
ening process which optimizes osmotic adjustment. However more
studies are required to test if there are other traits in combination
with osmotic adjustment that allow water stress tolerance without
penalization in tuber yield.
Acknowledgement
This research was conducted under the CGIAR Research Pro-
gram on Water, Land and Ecosystems. Authors thank Felipe de
Mendiburu for his support in the statistical analysis, and to Luis
Silva and Nikolai Alarcon for their assisstance in field assessments.
References
Blum, A., 2011. Drought resistance and its improvement. In: Blum, A. (Ed.), Plant
Breeding for Water-limited Environments. Springer, New York, pp. 53–152.
Bruce, J.A., Matthes, M.C., Napier, J.A., Pickett, J.A., 2007. Stressful ‘memories’ of
plants: evidence and possible mechanisms. Plant Science 173, 603–608.
FAO, 2012. Irrigation water management: irrigation methods. Annex 2 Infiltration
rate and infiltration test. http://www.fao.org/docrep/S8684E/s8684e0a.htm
Gutiérrez, R., Espinoza, J., Bonierbale, M., 2007. UNICA: variedad Peruana para mer-
cado fresco y papa frita con tolerancia y resistencia para condiciones climáticas
adversas. Revista Latinoamericana de la Papa 14 (1), 41–50.
Haverkort, A.J., Verhagen, A., Grashoff, C., Uithol, P.W.J., 2004. Potato-zoning:
a decision support system on expanding the potato industry through agro-
ecological zoning using the LINTUL simulation approach. In: MacKerron, D.K.L.,
Haverkort, A.J. (Eds.), Decision Support Systems in Potato Production, Bring-
ing Models to Practice. Wageningen Academic Publishers, The Netherlands,
pp. 28–44.
Hijmans, R.J., 2003. The effect of climate change on global potato production. Amer-
ican Journal of Potato Research 80 (4), 271–279.
Jefferies, R.A., 1995. Physiology of crop response to drought. In: Haverkort, A.J.,
MacKerron, D.K.L. (Eds.), Potato Ecology and Modeling of Crops under Condi-
tions Limiting Growth. Wageningen Academic Publishers, The Netherlands, pp.
61–74.
70 W. Yactayo et al. / Agricultural Water Management 123 (2013) 65–70
Jensen, C.R., Battilani, A., Plauborg, F., Psarras, G., Chartzoulakis, K., Janowiak, F., Sti-
kic, R., Jovanovic, Z., Li, G.T., Qi, X.B., Liu, F.L., Jacobsen, S.E., Andersen, M.N., 2010.
Deficit irrigation based on drought tolerance and root signalling in potatoes and
tomatoes. Agricultural Water Management 98 (3), 403–413.
Jovanovic, Z., Stikic, R., Vucelic-Radovic, B., Paukovic, M., Brocic, Z., Matovic, G., Rov-
canin, S., Mojevic, M., 2010. Partial root-zone drying increases WUE, N and
antioxidant content in field potatoes. European Journal of Agronomy 33 (2),
124–131.
Kang, S., Zhang, J., 2004. Controlled alternate partial root-zone irrigation: its
physiological consequences and impact on water use efficiency. Journal of
Experimental Botany 55, 2437–2446.
Kundzewicz, Z.W., Mata, L.J., Arnell, N.W., Döll, P., Kabat, P., Jiménez, B., Miller,
K.A., Oki, T., Sen, Z., Shiklomanov, I.A., 2007. Freshwater resources and their
management. In: Parry, M.L., Canziani, O.F., Palutikof, J.P., van der Linden, P.J.,
Hanson, C.E. (Eds.), Climate Change 2007: Impacts, Adaptation and Vulnerability.
Contribution of Working Group II to the Fourth Assessment Report of the Inter-
governmental Panel on Climate Change. Cambridge University Press, Cambridge,
UK, pp. 173–210.
Larcher, W., 2003. Physiological Plant Ecology, Ecophysiology and Stress Physiology
of Functional Groups. Berlin, Springer-Verlag, 513 pp.
Liu, F.L., Shahnazari, A., Andersen, M.N., Jacobsen, S.E., Jensen, C.R., 2006a. Effects
of deficit irrigation (DI) and partial root drying (PRD) on gas exchange, biomass
partitioning, and water use efficiency in potato. Scientia Horticulturae 109 (2),
113–117.
Liu, F.L., Shahnazari, A., Andersen, M.N., Jacobsen, S.E., Jensen, C.R., 2006b. Physiolog-
ical responses of potato (Solanum tuberosum L.) to partial root-zone drying: ABA
signalling, leaf gas exchange, and water use efficiency. Journal of Experimental
Botany 57 (14), 3727–3735.
Morgan, J.M., 1983. Osmoregulation as a selection criterion for drought
tolerance in wheat. Australian Journal of Agricultural Research 34 (6),
607–614.
Morgan, J.M., 1992. Osmotic components and properties associated with genotypic
differences in osmoregulation in wheat. Aust. J. Plant Physiol. 19, 67–76.
Morgan, P.W., Drew, M.C., 1997. Ethylene and plant responses to stress. Physiologia
Plantarum 100, 620–630.
Naylor, R.L., 2009. Managing food production systems for resilience. In: Chapin
III, S., Kofinas, G.P., Folke, C. (Eds.), Principles of Ecosystems Stewardship,
Resilience-based Natural Resources Management in a Changing World. Springer,
USA, pp. 259–280.
Rojas, G., Posadas, A., Quiroz, R., Holle, M., Málaga, M., 2007. Secado parcial de las
raíces: una promisoria técnica de riego en papa (Solanum tuberosum L.). Zonas
Áridas 11 (1), 206–218.
Saeed, H., Grove, I.G., Kettlewell, P.S., Hall, N.W., 2008. Potential of partial rootzone
drying as an alternative irrigation technique for potatoes (Solanum tuberosum).
Annals of Applied Biology 152 (1), 71–80.
Shahnazari, A., Liu, F.L., Andersen, M.N., Jacobsen, S.E., Jensen, C.R., 2007. Effects of
partial root-zone drying on yield, tuber size and water use efficiency in potato
under field conditions. Field Crops Research 100 (1), 117–124.
Shahnazari, A., Ahmadi, S.H., Laerke, P.E., Liu, F., Plauborg, F., Jacobsen, S.E., Jensen,
C.R., Andersen, M.N., 2008. Nitrogen dynamics in the soil–plant system under
deficit and partial root-zone drying irrigation strategies in potatoes. European
Journal of Agronomy 28 (2), 65–73.
Sharp, R.E., 2002. Interaction with ethylene: changing views on the role of abscisic
acid in root and shoot growth responses to water stress. Plant, Cell and Environ-
ment 25, 211–222.
Tardieu, F., 2012. Any trait or trait-related allele can confer drought tolerance: just
design the right drought scenario. Journal of Experimental Botany 63, 25–31.
Thiele, G., Theisen, K., Bonierbale, M., Walker, T., 2010. Targeting the poor and
hungry with potato science. Potato Journal 37, 75–86.
Thomas, H., Howarth, C., 2000. Five ways to stay green. Journal of Experimental
Botany 51, 329–337.
Turner, N.C., 1981. Techniques and experimental approaches for the measurement
of plant water status. Plant Soil 58, 339–366.
van Loon, C.D., 1981. The effect of water stress on potato growth, development, and
yield. American Potato Journal 58 (1), 51–69.
Wolfinger, R.D., Chang, M., 1998. Comparing the SAS GLM and MIXED Procedures
for Repeated Measures. In: SUGI Proceedings, Cary, North Carolina, USA.
Xie, K., Wang, X.-X., Zhang, R., Gong, X., Zhang, S., Mares, V., Gavilán, C., Posadas,
A., Quiroz, R., 2012. Partial root-zone drying irrigation and water utilization
efficiency by the potato crop in semi-arid regions in China. Scientia Horticulturae
134, 20–25.
Xu, H.L., Qin, F.F., Xu, Q.C., Tan, J.Y., Liu, G.M., 2011. Applications of xerophytophysi-
ology in plant production – the potato crop improved by partial root zone drying
of early season but not whole season. Scientia Horticulturae 129 (4), 528–534.