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Applying hydraulic lift in an agroecosystem: Forage plants with shoots

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Plant Soil
DOI 10.1007/s11104-010-0581-1
REGULAR ARTICLE
Applying hydraulic lift in an agroecosystem: forage
plants with shoots removed supply water to neighboring
vegetable crops
Nobuhito Sekiya & Hideki Araki & Katsuya Yano
Received: 6 June 2010 / Accepted: 14 September 2010
# Springer Science+Business Media B.V. 2010
Abstract When a plant encounters spatially heteroge-
neous soil moisture within its root system, usually drier
surface and moister subsurface soils, water can move
between these layers through the root system, a plant
process known as hydraulic lift or redistribution. The
water thus transferred is available not only for the plant
itself but also for its neighbors. We examined applica-
tion of this process as a possible biological irrigation
tool. As ‘donors’, we used perennial forage plants with
Responsible Editor: Hans Lambers.
Electronic supplementary material The online version of this
article (doi:10.1007/s11104-010-0581-1) contains
supplementary material, which is available to authorized users.
N. Sekiya (*)
Japan International Cooperation Agency (JICA),
JICA Tanzania Office, P. O. Box 9450 Dar es Salaam,
Tanzania
e-mail: kapinivilage@yahoo.co.jp
H. Araki
Experimental Farm, Faculty of Agriculture,
Yamaguchi University,
Yoshida,
Yamaguchi 753-8515, Japan
K. Yano
Laboratory of Crop Science, Graduate School
of Bioagricultural Sciences, Nagoya University,
Chikusa,
Nagoya, Aichi 464-8601, Japan
their shoots removed to minimize the effect of light-
interception by them on the ‘receiver’ plants growing
alongside them. In a horizontally split-root experiment,
where an upper container was filled with sand and a
lower one with water, superior donor species could
maintain the upper sand in a fully hydrated condition for
several weeks, increasing stomatal conductance in the
receivers. The effects were also confirmed in a water-
limited agricultural field, as significant differences were
found in canopy temperature and yield in neighboring
crop plants in the presence or absence of donor root
systems. These results suggest that deep-rooting associ-
ate plants with their shoots removed function as an
irrigation tool and improve crop production in water-
scarce environments.
Keywords Deep roots . Drought stress . Facilitation .
Intercropping . Positive plant–plant interaction .
Sustainable agriculture
Introduction
Plant roots absorb water from the soil but occasionally
release it back (Caldwell et al. 1998; Jackson et al.
2000). Water efflux from roots is regulated by the same
mechanisms as water influx into roots. Water influx
takes place along a water potential gradient from the
soil to the root xylem, and this gradient is generated by
transpirational water loss from the leaves. At night,

reduced transpiration allows xylem water potential to
rise (become less negative) and can reverse the
direction of the water potential gradient. Under these
conditions, water may exit the roots to the soil along a
reversed water potential gradient.
In deep-rooting systems that penetrate dry, upper
soil layers to reach wet, deeper soil layers, reduced
transpiration induces water efflux, mainly from the
shallow roots to the dry soil (Caldwell et al. 1998;
Jackson et al. 2000). This process drives an upward
water movement within the root system that has the
effect of rendering the soil water status within the
whole root zone more homogeneous. The process is
termed hydraulic lift as water is often lifted from
deeper to shallower soils through the root system
(Caldwell and Richards 1989; Richards and Caldwell
1987). In fact, water will actually move from one
body of soil to another, just so long as a water
potential gradient exists between them so the water
movement is not necessarily in an upward direction
(Hultine et al. 2004; Oliveira et al. 2005; Schulze et
al. 1998; Smart et al. 2005; Warren et al. 2007).
Therefore, in a more expanded concept, hydraulic lift
is sometimes termed hydraulic redistribution (Burgess
et al. 1998, 2000).
Water efflux was first observed about 80 years ago
in a split-root experiment in which the root portion in
a dry soil exuded water while the roots of the other
portion were in a wet soil (Braezeale 1930). Subse-
quently, new techniques, using a stable isotope
method in particular, made it possible to detect water
efflux and hydraulic lift even under field conditions
(Burgess et al. 2000; Dawson 1993, 1996). As a
result, examples have been noted in many species at
not only laboratory (Hawkins et al. 2009; Leffler et al.
2004; McMichael and Lascano 2010; Valenzuela-
Estrada et al. 2009; Wan et al. 2000; Xu and Bland
1993) but also field scales (Caldwell et al. 1998;
Espeleta et al. 2004; Jackson et al. 2000; Liste and
White 2008). The stable isotope method further
revealed that the hydraulically lifted groundwater is
even transferred to neighboring plants as if the lifting
plant irrigates its neighbors with groundwater (Armas
et al. 2010; Dawson 1993; Ludwig et al. 2003;
Moreira et al. 2003). Based on these reports, a new
idea emerged that roots could function as an irrigation
system if the hydraulic lift was managed appropriately
(Liste and White 2008; Sekiya and Yano 2004). If this
can be arranged in an agricultural field, a novel and
Plant Soil
practical benefit arises based on the hydraulic lift/
redistribution process.
Therefore, a series of experiments was conducted
both in the laboratory and the field to examine the
hypothesis that the process of hydraulic lift can be
applicable to irrigation of neighboring crops. We first
evaluated water lifting capabilities of donor plants in
a horizontally split-root system to determine the
potential water supply. Next, using a similar but
larger system, we assessed effects of water supply
provided by donor plants on water relations of
receiver crops. Finally, the effects were tested in a
water-limited agricultural field.
Materials and methods
Selection of plant species
Attempts were made to find effective donor species
having high water-lifting capability. We considered
the following two criteria as prerequisites for candi-
date selection. 1) Deep rooting: hydraulic lift is more
likely to occur with a deep-rooting species. 2) Forage
species: Removal of leaves by cutting the shoot is
expected to minimize the adverse effect of light-
interception on the receiver plants growing alongside
them. For this purpose, forage plants seemed suitable
considering their tolerance to shoot removal (grazing).
The following species were investigated: guinea grass
(Panicum maximum), tall fescue (Festuca arundina-
cea), white clover (Trifolium repens), smooth brome-
grass (Bromus inermis) and sun hemp (Crotalaria
juncea). Pigeon pea (Cajanus cajan) was also
compared with these species because it was previous-
ly reported to have water-lifting capability (Sekiya
and Yano 2004).
Comparison of water-lifting capabilities
Split-root experiment Two plastic containers (0.32 m
long × 0.1 m wide × 0.1 m high) were placed one on top
of the other (Fig. 1a). A rectangular hole (0.1 m long ×
0.025 m wide) was first cut in the bottom of the upper
container near one end. This hole was covered with
0.15 mm nylon mesh to contain the rooting medium
while still allowing root penetration. The upper
container held 3.5 kg of sand while the lower one
was filled with 2.0 kg of water. There was an air gap
Plant Soil
Fig. 1 Comparison of water lifting capability. a A schematic of
the split-root container. b Sand surfaces grown without associated
donor plants (Cont) and with Cajanus cajan (Cc), Crotalaria
juncea (Cj) and Panicum maximum (Pm). c The amount of water
lifted from the lower container per unit of water evaporated from
the upper one (HL Eup−1) in split-containers with Cc, Cj, Bromus
inermis (Bi), Trifolium repens (Tr), Festuca arundinacea (Fa)
and Pm. Each bar represents mean ± S.E. (n=5). The letters
above the bars represent significant differences at the 5% level as
analyzed by Sheffe’s F test
between the containers. Between 10 and 20 germinated
seeds each of P. maximum, F. arundinacea, T. repens,
B. inermis, C. juncea and C. cajan were sown in the
upper container just above the hole. The same setup
but without plants served as a control. Each test
donor species and the control were replicated five
times. Water was supplied at 2- to 3-day intervals
to the upper container to maintain field capacity
(defined as the medium’s water content 24 h after
saturation) and the amount of water in the lower
container was maintained at 2.0 kg. Between 1 and
3 weeks after sowing, seedlings were thinned to
leave several uniform and healthy plants. To avoid
too marked differences in rooting volume between
the donor species, C. juncea and C. cajan seedlings
were thinned to 4 individuals, P. maximum to 5, B.
inermis to 7, F. arundinacea to 12 and T. repens to
10. A commercially available liquid fertilizer (Hypo-
nex 6-6-6; Hyponex Japan, Osaka, Japan) diluted
100 times with deionized water was applied to the
upper container twice and a thousand-times solution
to the lower one three times during growth. By 10 to
13 weeks after sowing, roots were well developed in
the lower container for all donor species. The shoots
were then cut at the base and the exposed surfaces
covered with Parafilm. To measure the amount of
water lifted from the lower to the upper container
(HL), the water supply to the upper container was
withheld. HL was calculated as
HL ¼ Llow  Elow
where Llow was water loss from the lower container
measured by weighing the container every day, and
Elow was evaporation from the lower container
estimated by weighing the lower container of the
controls every day. Evaporation from the upper
container (Eup) was also calculated as
Eup ¼ Lup þ HL
where Lup was water loss from the upper container
measured by weighing the container every day. The
amount of water lifted relative to the amount
evaporating from the upper container (HL Eup−1)
was calculated by dividing HL by Eup to compare
water-lifting capability among the plant species.
Water was supplied to the lower containers to
maintain their initial 2.0 kg volume after every
measurement. The upper containers were supplied
with water to field capacity when their water
contents fell below 50% of field capacity to avoid
root desiccation and death. At 122 days after sowing,
the roots in the top containers were gently washed
free of sand under running water above a 2-mm
sieve. The collected roots were preserved in FAA
(formalin : acetic acid : 70% ethanol = 1 : 1 : 18, v/v/v)

for root length measurement. The roots were then
stained with 0.2% (w/v) methylene blue for 48 h and
spread on a transparent sheet, without overlap. Digi-
tized images were produced using a scanner with
resolution of 300 dpi and an output format of 256 grey-
scales. The root length was determined using a macro-
program developed by Kimura et al. (1999) on NIH
Image software (version 1.60). Water-lifting capability
per unit of root length in the upper containers was
calculated by diving HL Eup−1 by root length.
Effect of hydraulic lift on neighboring crop growth
A similar but larger split-root system was employed
using a container (0.4 m long × 0.32 m wide × 0.25 m
deep) as shown in Online Resource 1. This was placed
on top of a smaller container (15 L). A rectangular hole
(0.3 m long × 0.02 m wide) was cut in the bottom of
the upper container and covered with nylon mesh as
before. The upper container was filled with 30 L of
sand while the lower one was filled with 15 L of water.
Again, there was an air gap between the two containers.
The sand was mixed with 1.10 g kg−1 of compound
fertilizer (N : P2O5 : K2O=0.08 : 0.08 : 0.08 g g−1),
0.5 g kg−1 of compound trace element fertilizer (MgO :
MnO : B2O3 =0.14 : 0.003 : 0.003 g g−1) and
0.29 g kg−1 of slaked lime. Twenty germinated seeds
each of T. repens, C. juncea, F. arundinacea and B.
inermis were sown in the upper containers just above
the rectangular hole. The same setup but without donor
plants served as a control. Each species and its control
were replicated four times. Water was supplied at 2- to
3-day intervals to maintain field capacity in the upper
containers. At 99 days after sowing, the donor plant
roots started to grow into the lower containers and the
water there was then aerated to maintain good aerobic
conditions. Water was supplied to the lower containers
at 2- to 3-day intervals to maintain the initial amount. At
127 days after sowing, the donor shoots were cut at the
base and the exposed surfaces covered with Parafilm. At
153 days from sowing, cucumber (Cucumis sativus L.)
seeds were sown adjacent to the donor plants at
50×50 mm spacing. Before sowing, 5 L of nutrient
solution (containing 3.90 mg N, 1.80 mg P2O5, 6.10 mg
K2O, 3.45 mg CaO, 0.90 mg MgO, 0.023 mg MnO,
0.023 mg B2O3, 0.04 mg Fe, 0.45 μg Cu, 1.35 μg Zn
and 0.45 μg Mo) was applied to each upper container to
field capacity, and further watering of the upper
Plant Soil
container was withheld. Volumetric soil moisture con-
tent (m3 m−3) was measured using a soil moisture sensor
(ECH2O EC-5; Decagon Devices, WA, USA) buried
0.2 m from the donor plants and 50 mm below the soil
surface in each upper container. Data readings were
made using a data logger (Em 50; Decagon Devices,
WA, USA) 5, 9, 13, 17, 24, 27 and 30 days from
sowing C. sativus. The sensor was calibrated with the
same sand as used in this experiment adjusted at 0, 0.06
and 0.10 m3 m−3 soil moisture content. Stomatal
conductance (gs) was measured on the abaxial surface
of the leaf using a hand-held diffusion porometer
(Porometer AP4; Delta-T, Cambridge, UK) at 22, 29
and 34 days from sowing. C. sativus plants were
harvested 35 days after sowing. The plants were dried
at 80°C for 48 h and dry weights were measured.
Effect of hydraulic lift on crop production under field
conditions
The field experiment was conducted at the Jinzai
Sand Dune Experimental Field of Shimane University
(N 35° 20′ 6, E 132° 41′ 3) between March 2007 and
October 2007. Seedlings of P. maximum, F. arundi-
nacea, T. repens, B. inermis, C. juncea and C. cajan
were established in PVC tubes (0.5 m diameter × 1 m
long) 9 weeks before the start of the experiment. Ten
seedlings of each donor species were transplanted
along the center line of each plot (3 m long × 1.5 m
wide) at 0.15-m intervals on 11 May, 2007 as shown
in Online Resource 2. Iron sheets were buried
vertically to a depth of 0.5 m to prevent root
interactions between plots. At 80 days after trans-
planting, transparent PVC covers were constructed
over the experimental plots to eliminate rainfall and
water was supplied directly to the plants by drip
irrigation. At 110 days after transplanting, the shoots
were cut at a height of 0.1 to 0.3 m from the soil
surface and the drip irrigation was replaced with
sprinklers to supply water evenly over the whole plot.
Next, 250 g of compound fertilizers (N : P2O5 :
K2O=0.14 : 0.26 : 0.14 g g−1) and 250 g of
magnesium lime were spread over each plot. At
130 days after transplanting, komatsuna (Brassica
rapa var. peruviridis) seeds were sown at 0.1×0.05 m
spacing in each plot as shown in Online Resource 2.
At the two- to three-leaf stage, the B. rapa seedlings
were thinned to leave a single plant at each sowing
point. Then, the roots on one side of the donor species
Plant Soil
were cut by inserting an iron sheet with a sharp edge
to 0.5 m depth along the center line while the roots on
the other side were left intact as shown in Online
Resource 3. At the four- to five-leaf stage, water was
supplied to the B. rapa plants only when most of the
plants had wilted. Images of the B. rapa plants were
captured using thermography (Thermo Tracer
TH7800N Fusion; NEC-Sanei, Osaka, Japan). At
45 days after sowing, the B. rapa plants growing
within 0.3 m of the center line were harvested and
their fresh matter weights were measured.
Statistical analyses
The water lifted per unit of water evaporated (HL Eup−1
or water-lifting capability), root length and the
water-lifting capability per unit of root length were
analyzed by Sheffe’s F test, soil moisture content and
gs by a fixed-model two-way analysis of variance
(ANOVA) in which sources of variation were the
measurement date, the donor species and their
interactions, C. sativus dry weight by Tukey-
Kramer test and B. rapa fresh weight by a mixed-
model ANOVA (strip-plot design) with the donor
species as a horizontal factor and the root cutting
treatment as a vertical factor. Correlations between
water-lifting capability and root length were ana-
lyzed by Pearson correlation coefficient test.
Results
Comparison of water-lifting capabilities
The water-lifting capabilities of selected plant species
were compared in a horizontally split-root container
(Fig. 1a). After withholding water from the upper
container, the split-container without roots (control)
rapidly lost water from the upper container due to
evaporation, as could be seen by the pale color of the
sand (Fig. 1b). In contrast, the split containers with roots
maintained substantial water content, depending on the
species (Fig. 1b). For instance, P. maximum kept all the
sand’s surface wet until the end of the experiment, while
C. juncea partially wetted it. C. cajan roots, however,
kept little water on the sand’s surface.
To determine water-lifting capability, the amount
of water lifted from the lower container per unit of
water evaporated from the upper one (HL Eup−1) was
calculated based on gravimetric measurements of the
water balance in the split-root container (Fig. 1c). The
values of P. maximum, F. arundinacea and T. repens
ranged between 0.913 and 0.996 g g−1, indicating that
the amount of water evaporating from the upper
container was almost fully replaced by the water lifted
from the lower one. B. inermis, C. juncea and C.
cajan had the small values compared with those
species with high level of water-lifting. The value for
C. cajan, which was the smallest among the tested
species, was 0.148 g g−1, showing that only 15% of
the water evaporating from the upper container was
lifted from the lower one. Surprisingly, roots of
P. maximum and F. arundinacea maintained high
levels of water lifting even after 1 month and the total
amount of water lifted was far greater than expected.
Root length and its relationship with water-lifting
capability were analyzed to investigate causes of
interspecific differences in water-lifting capability
(Table 1). Root length ranged between 18 and
524 m container−1 among the species. The multiple
comparisons indicate that the species can be divided
into two groups; one with longer roots (P. maximum,
F. arundinacea and B. inermis) and the other with
shorter ones (T. repens, C. juncea and C. cajan). The
correlation coefficient test was carried out to analyze
the relationship between the water-lifting capability
(HL Eup−1) and root length. The results showed that
there was no significant correlation between the two
parameters (R=0.280, P=0.157). To further investi-
gate causes of interspecific differences in water-lifting
capability, the water-lifting capability per unit of root
length was calculated. The multiple comparisons
indicate that there was no significant difference in
this value among C. cajan, C. juncea, B. inermis, F.
arundinacea and P. maximum while the value of T.
repens was almost 23 times larger than those of the
others. Thus, the relationship between the water-
lifting capability and root length was tested again
after excluding the data of T. repens. The results
showed that the capability was significantly correlated
with root length (R=0.607, P=0.003). These results
suggest that while the interspecific differences in
water-lifting capability was caused by root length or
root length density among C. cajan, C. juncea, B.
inermis, F. arundinacea and P. maximum, the high
level of water-lifting in T. repens was due to the high
level of water-lifting capability per unit of root length
or water permeability of roots.
 Plant Soil

Table 1 Relationship between root length and hydraulic lift

Cc Cj Bi Tr Fa Pm

Root length (m container−1) 132±29b 87±7b 498±45a 18±4b 394±35a 524±10a

Water-lifting capability per root legnth (g g−1 m−110−3) 3.53±3.1b 3.67±1.5b 1.24±0.3b 60.29±12.6a 2.54±0.2b 1.89±0.04b

Root length of Cajanus cajan (Cc), Crotalaria juncea (Cj), Bromus inermis (Bi), Trifolium repens (Tr), Festuca arundinacea (Fa) and
Panicum maximum (Pm) were determined using a macro-program developed by Kimura et al. (1999) on NIH Image software (version
1.60). Water-lifting capability was determined as the amount of water lifted from the lower container per unit of water evaporated from
the upper one in split-containers as shown in Fig. 1c. Each value represents mean ± S.E. (n=5). The values with different letters in a
line are significantly different at the 5% level as analyzed by Sheffe’s F test

Effect of hydraulic lift on neighboring crop growth
The effect of hydraulic lift on neighboring plants was
assessed in terms of their water status and growth in a
similar but larger split-root container (Online Re-
source 1). After saturating the sand in the upper layer,
C. sativus seeds were sown next to the water-lifting
plants, and water was withheld from the upper layer at
sowing. Five days after sowing, all containers still
maintained similar moisture levels ranging between
0.081 and 0.106 m3 m−3 (Fig. 2a). While the value in
the control declined due to evapotranspiration after-
wards, they remained relatively high in the presence
of the donor plants. For instance, at 17 days after
sowing, the soil moisture content of the control was
0.015 m3 m−3, and those with the donor plants ranged
between 0.054 and 0.076 m3 m−3. The soil moisture
contents were shown by a two-way ANOVA to be
significantly different among the measuring dates
(P<0.001) and the donor species (P<0.001). There
was no significant interaction between the two effects
(P=0.994). These results suggest that the donor roots
can maintain soil moisture content at relatively high
level by water-lifting. As a result, the C. sativus plants
had wide, turgid leaves in the presence of donor roots
while in their absence the leaves wilted (Fig. 2b).
During the C. sativus growth period, the stomatal
conductance (gs) of the leaves was also measured
three times (Fig. 2c). At the first measurement, gs of
C. sativus leaves from all containers were relatively
low due to low light intensity. At the second and third
measurements, gs became higher in the presence of
the donor plants, though the extent in the presence of
F. arundinacea was lower than with T. repens, B.
inermis or C. juncea. In contrast, gs in the control
remained low. gs values were shown by a two-way
ANOVA to be significantly different among the
measuring dates (P=0.001) and the donor species
(P< 0.001). There was no significant interaction
between the two effects (P=0.419). These results
suggest that water lifted by the donor plants can
increase the gs of neighboring C. sativus plants. The
C. sativus plants irrigated by T. repens, B. inermis and
C. juncea had significantly higher gs than those in the
control, resulting in greater biomass accumulation
(Fig. 2d).
Effect of hydraulic lift on crop production under field
conditions
The effect of hydraulic lift on crop production was
tested in a water-limited field situation (Online
Resource 2). The plants of each donor species were
established along the center line in each experimental
plot, and the roots on one side of the donor plants
were severed to prevent water supply through
hydraulic lift while those on the other side were left
intact (Online Resource 3). As expected, the presence
of intact donor roots allowed B. rapa to maintain
fresh and turgid leaves while in the absence of intact
roots, the B. rapa leaves wilted due to water deficit
(Fig. 3).
Thermal images of B. rapa leaves captured during
the growth period show that leaf temperatures of B.
rapa plants associated with the intact donor roots
were lower than those without (Fig. 4). The differ-
ences in leaf temperature between the intact- and
cut-treatments were particularly distinct in P. maxi-
mum, F. arundinacea, C. juncea and B. inermis. Leaf
temperature is lowered by transpiration, so reduced
leaf temperature demonstrates that these B. rapa
plants associated with intact donor roots had en-
hanced transpiration. This also implies increased
stomatal opening and hence increased photosynthesis.
The fresh matter weights of B. rapa shoots
associated with intact donor roots were shown by a
Plant Soil
Fig. 2 Effects of hydraulic lift on neighboring crop growth. a
Soil moisture content of shallow soils (50 mm below soil
surface) grown without associated donor plants (Cont) and with
Festuca arundinacea (Fa), Crotalaria juncea (Cj), Bromus
inermis (Bi) and Trifolium repens (Tr) measured 5, 9, 13, 17,
24, 27 and 30 days after sowing Cucumis sativus. Each data
point represents mean ± S.E. (n=4). A fixed model two-way
analysis of variance showed that soil moisture content was
significantly different among the measuring dates (P<0.001)
and the donor species (P<0.001). There was no significant
interaction between the two effects (P=0.994). b Leaves of C.
mixed-model ANOVA (strip-plot design) to be sig-
nificantly larger than those without (P= 0.049)
(Fig. 5). This result suggests that the donor plants
supplied hydraulically lifted water to B. rapa plants
through their intact roots. The mixed-model ANOVA
also showed that the fresh matter weights of B. rapa
shoots were significantly different among the donor
species (P<0.001), demonstrating an interspecific
difference in water-supply capability. The increase in
B. rapa growth was particularly high in association
with P. maximum, F. arundinacea and C. juncea. In
the cut-treatment, there was also a difference between
sativus plants. c Stomatal conductance (gs) of the abaxial
surfaces of the C. sativus leaves measured 22, 29 and 34 days
after sowing. Each data point represents mean ± S.E. (n=4). A
fixed model two-way analysis of variance showed that gs was
significantly different among the measuring dates (P=0.001)
and the donor species (P<0.001). There was no significant
interaction between the two effects (P=0.419). d Dry matter
weight of C. sativus plants. Each bar represents mean ± S.E.
(n=4). The letters above the bars represent significant differ-
ences at the 5% level analyzed by Tukey-Kramer test
the control (the absence of roots) and the donor
species (the presence of cut roots). This implies that
the presence of roots, regardless of being dead or
alive, increased B. rapa growth. There was no
significant interaction between the cut-treatment and
the donor species (P=0.153).
Discussion
The experiments conducted in the present study
clearly demonstrated that the process of hydraulic lift
 Plant Soil

can be applicable for irrigation of neighboring crops.

Fig. 3 Brassica rapa plants in Panicum maximum plot with
intact P. maximum roots (Intact) and with cut P. maximum roots
(Cut) under field conditions
The split-root experiment revealed that all the plant
species examined in this study were capable of lifting
a substantial amount of water (Fig. 1c). The lifted
water was further shown to improve the water status
of neighboring crops, and as a result, increase their
shoot dry matter weight (Fig. 2d). The positive effects
of donor roots on the water status (Figs. 3 and 4) and
plant growth (Fig. 5) of neighboring vegetables were
confirmed even in a water-limited agricultural field.
Therefore, we conclude that perennial forage plants
are able to function as an effective biological
irrigation system.
The large split-root experiment demonstrated that
the donor roots maintain high levels of soil moisture
content without additional water supply as a conse-
quence of hydraulic lift while the control lost

Fig. 4 Thermal images of

Brassica rapa plants grown
under field conditions with-
out associated donor plants
(Cont) and with Cajanus
cajan (Cc), Trifolium repens
(Tr), Bromus inermis (Bi),
Crotalaria juncea (Cj),
Festuca arundinacea (Fa)
and Panicum maximum
(Pm). The donor roots were
left intact on one side
(Intact) and cut on the
other (Cut)
Plant Soil
Fig. 5 Fresh matter weight of Brassica rapa plants grown
under field conditions without associated donor plants (Cont)
and with Cajanus cajan (Cc), Trifolium repens (Tr), Bromus
inermis (Bi), Crotalaria juncea (Cj), Festuca arundinacea (Fa)
and Panicum maximum (Pm). Plants with intact donor roots
(gray bars) and with cut donor roots (white bars) are presented
separately. Each bar represents mean ± S.E. (n=4). A mixed-
model analysis of variance (strip-plot design) showed that fresh
matter weights were significantly different among the donor
species (P < 0.001) and between the root cut treatments
(P=0.049). There was no significant interaction between the
two effects (P=0.153)
substantial amount of water due to evapotranspiration
(Fig. 2a). In contrast, no distinct differences in soil
moisture content were observed between the donor
plots and the control ones in the field experiment
(data not shown). We would attribute this contradic-
tion to a position of sensor in the field experiment.
The thermal images suggest that the effect of donor
roots reached only 0.2 m to 0.3 m from the center line
in each plot (Fig. 4). As a result, the growth
enhancement was pronounced only in those B. rapa
plants grown within 0.3 m from the donors (Fig. 5). In
this experiment, however, a soil moisture sensor
(Delta-T Devices, Cambridge, UK) was buried
0.6 m from the donors. The sensor is assumed to
have measured moisture content of soils away from
which the donor roots could function. Monitoring of
soil moisture change will allow us to predict effects of
hydraulic lift at different scales and environments
(Amenu and Kumar 2008; Araki et al. 2008; Doussan
et al. 2006; Siqueira et al. 2008). Therefore, the extent
to which the effect of hydraulic lift reaches should be
fully investigated to precisely understand soil mois-
ture dynamics under field conditions.
Previously, hydraulic lift was observed mainly in
the laboratory (Hawkins et al. 2009; Leffler et al.
2004; McMichael and Lascano 2010; Valenzuela-
Estrada et al. 2009; Wan et al. 2000; Xu and Bland
1993) and in natural ecosystems (Caldwell et al.
1998; Espeleta et al. 2004; Jackson et al. 2000; Liste
and White 2008). Little attention has been paid to its
role in agroecosystems except in a few reports (Kizito
et al. 2007; Sekiya and Yano 2004). This study
confirms not only that hydraulic lift can occur in an
agricultural situation but also that its influence on
crop production may be unexpectedly high. Several
field studies demonstrated that woody plants supply
hydraulically lifted water to understory grasses
(Brooks et al. 2006; Dawson 1993; Ludwig et al.
2003, 2004; Moreira et al. 2003). However, few of
these studies observed a positive effect of the water
supply on understory plant production except for
Dawson (1993), who found that understory grasses
had higher plant water potential and stomatal con-
ductance. These reports argue that the positive effect
of water supply is overwhelmed by a negative effect
of soil water competition between woody plants and
understory grasses (Ludwig et al. 2003, 2004) or that
the positive effect is only minimal (Brooks et al.
2006; Moreira et al. 2003). As we proposed previ-
ously (Sekiya and Yano 2004), defoliation and
pruning may enhance the water supply capability of
donor plants by reducing their transpiration. Leffler et
al. (2005) demonstrated that hydraulic lift occurs even
in senescent or shoot-removed plant roots as long as
they remain intact and hydrated. Forage plants with
their shoots removed in the present study might have
also enhanced neighboring vegetable growth through
a reduction in soil water competition and/or a
continued water supply even during daylight. A
further study is needed to reveal and quantify
contribution of each process to the enhancement of
neighboring vegetable growth in agroecosystems.
P. maximum and F. arundinacea displayed partic-
ularly high performances for irrigation under field
conditions (Fig. 5). Previously, biological irrigation
was demonstrated only with C. cajan in an earlier
study of ours (Sekiya and Yano 2004). Using the
hydrogen stable isotope method, it was revealed that
C. cajan hydraulically lifted groundwater and sup-
plied it to neighboring Zea mays plants in a field in
Zambia. The performance was shown to be enhanced
further by shading the shoots of C. cajan in a
controlled environment. The present study revealed
that forage plants possess higher water-lifting capa-
bilities than C. cajan (Fig. 1c). The superiority of
forage plants over C. cajan was also demonstrated in
water-supply capability (Figs. 2d and 5).
Interspecific differences in hydraulic lift are con-
sidered to be caused by differences in root length

density of donor plants in most cases (Table 1). Araki
et al. (2008) reported that water efflux from roots
increases with an increase in root length density. Also
indicated was that a sandy soil such as that in this
study reduces water efflux due to low soil hydraulic
conductivity. As indicated by Hansen and Dickson
(1979), water transfer between plants can be facilitat-
ed by closely associated, intertwined root systems. By
having a high root length density, therefore, the high-
performance species might have increased root con-
tact with the vegetable crops as well as enhanced
water efflux, making it possible to transfer more water
to the vegetable crops with little influence of low
hydraulic conductivity of the sandy soils. It is
plausible, however, that the high-performance of
T. repens is attained with the large permeability of
roots to water efflux rather than the large root length
density (Table 1). A numerical model to simulate
hydraulic lift predicted that the root permeability also
greatly influences water flow rate from roots to soils
as was the case in T. repens (Siqueira et al. 2008).
Factors such as thickness of cortex layer are respon-
sible for the large root permeability to water influx
(Huang and Eissenstat 2000; Rieger and Litvin 1999).
The same factors may have also contributed to the
large water efflux in T. repens in the present study.
This unique feature of T. repens should be a subject of
further study.
Although not yet properly tested, an idea has been
around for some time that crop yield in water-scarce
environments could be improved if new varieties with
deeper roots were to be bred (Huang 2000; Kramer and
Boyer 1995). However, it should be noted that a deeper
(higher biomass) root system will likely be associated
with some depression of shoot growth (Ma et al. 2010;
Markesteijn and Poorter 2009). Hence, deeper rooting
may not necessarily result in improved crop yield or
aboveground biomass production under water-scarce
environments. Any biomass gains made by the deeper
rooting may be substantially used up underground. In
contrast, it is proposed here that the water-lifting plants
are charged the biomass cost of deep root development.
This strategy could allow shallower-rooting crop
species to avoid severely depressed shoot growth as a
result of soil-water deficit.
The proposed association of donor and receiver
crops lifts water back up into the crop root zones that
would otherwise be lost without being acquired by the
crops. This would make possible more efficient use of
Plant Soil
a limited and precious water resource in water-scarce
environments. The technique is also expected to
increase soil nutrient availability for crops in water-
scarce environments (Aanderud and Richards 2009;
Caldwell and Manwarning 1994; McCulley et al.
2004; Vetterlein and Marschner 1993). Additionally,
using leguminous plants as donors also improves the
nitrogen nutrient status of the crops, as has been noted
previously (Sekiya and Yano 2004). In conjunction
with other water-saving techniques such as mulching
and minimum tillage (Ali and Talukder 2008;
Stroosnijder 2009; Wang et al. 2007), this technique,
involving an association between donor and recipient
species, is expected to further improve crop produc-
tion in water-scarce agroecosystems.
Conclusion
The perennial forage plants, P. maximum and F.
arundinaceae in particular, were shown to function as
effective biological irrigation systems. This system
would allow shallow rooting crops to recover draining
water without having them deep-rooted, making
possible the efficient use of irrigation water in water-
scarce environments. Potentially, further improvement
in the efficiency of water use could be achieved by a
joint application of biological irrigation and other
water-saving techniques. Future work should aim at
developing such a comprehensive technique for water-
scarce agroecosystems.
Acknowledgements We thank Y. Mizumoto, K. Iwamoto and
T. Furukawa for their cooperation in data acquisition. We are
also grateful to F. Adachi, M. Kadowaki, N. Yasuda and
T. Yamane for their help with the field work. This research was
supported financially by the Industrial Technology Research
Grant Program in 2005 from the New Energy and Industrial
Technology Development Organization (NEDO) of Japan, and
in part by KAKENHI (17688015, 20380178).
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